St R E L I T Z I a 39 (2018)

S T R E L I T Z I A 39 (2018) a b S T R E L I T Z I A 39 (2018) S T R E L I T Z I A 39

A TAXONOMIC REVISION OF Calobota (FABACEAE, CROTALARIEAE)

James Stephen Boatwright, Patricia May Tilney and Ben-Erik van Wyk

Pretoria 2018 S T R E L I T Z I A

This series has replaced Memoirs of the Botanical Survey of South Africa and Annals of the Kirstenbosch Botanic Gardens, which the South African National Biodiversity Institute (SANBI) inherited from its predecessor organisations.

The plant genus Strelitzia occurs naturally in the eastern parts of southern Africa. It comprises three arborescent species, known as wild bananas, and two acaulescent species, known as crane flowers or bird- of-paradise flowers. The logo of SANBI is partly based on the striking inflorescence ofStrelitzia reginae, a native of the Eastern Cape and KwaZulu-Natal that has become a garden favourite worldwide. It symbol- ises the commitment of SANBI to champion the exploration, conservation, sustainable use, appreciation and enjoyment of South Africa’s exceptionally rich biodiversity for all people.

James Stephen Boatwright Department of Biodiversity and Conservation Biology, University of the Western Cape, Private Bag X17, Bellville 7535, Cape Town, South Africa; e-mail: [email protected].

Patricia May Tilney and Ben-Erik van Wyk Department of Botany and Plant Biotechnology, University of Johannesburg, P.O. Box 524, Auck- land Park 2006, Johannesburg, South Africa; e-mail: [email protected]; [email protected].

EDITOR: Alicia Grobler PROOFREADER: Yolande Steenkamp DESIGN & LAYOUT: Elizma Fouché

Citing this publication: BOATWRIGHT, J.S., TILNEY, P.M. & VAN WYK, B-E. 2018. A taxonomic revision of Calobota (Fabaceae, Crotalarieae). Strelitzia 39. South African National Biodiversity Institute, Pretoria.

ISBN: 978-1-928224-27-3

Obtainable from: SANBI Bookshop, Private Bag X101, Pretoria 0001, South Africa Tel.: +27 12 843 5000 E-mail: [email protected] Website: www.sanbi.org Printed by: Rand Data Forms (Pty) Ltd, P.O. Box 24194, Gezina, 0031 (Pretoria Office), tel. no.: +27 12 567 1564. Copyright © 2018 by South African National Biodiversity Institute (SANBI). All rights reserved. No part of this book may be reproduced in any form without written permission of the copyright owners. The views and opinions expressed do not necessarily reflect those of SANBI. The authors and publisher have made their best efforts to prepare this book, and make no representation or warranties of any kind with regard to the com- pleteness or accuracy of the contents herein. All images in this book have been reproduced with the knowledge and prior consent of the artists concerned and no responsibility is accepted by the publisher or printer for any infringe- ment of copyright or otherwise arising from the contents of this publication. Every effort has been made to ensure that the credits accurately comply with the information supplied by the authors. S T R E L I T Z I A 39 (2018) iii

Contents

Abstract...... iv Acknowledgements...... v Introduction...... 1 Ethnobotany and chemistry...... 3 Phylogenetic relationships ...... 5 Horticultural potential...... 6 Materials and methods ...... 7 Vegetative morphology and anatomy...... 8 Reproductive morphology and anatomy...... 13 Taxonomic treatment...... 19 Calobota Eckl. & Zeyh...... 19 Key to the species of Calobota ...... 21 1. Calobota acanthoclada (Dinter) Boatwr. & B.-E.van Wyk...... 22 2. Calobota angustifolia (E.Mey.) Boatwr. & B.-E.van Wyk...... 25 3. Calobota cinerea (E.Mey.) Boatwr. & B.-E.van Wyk ...... 33 4. Calobota cuspidosa (Burch.) Boatwr. & B.-E.van Wyk...... 37 5. Calobota cytisoides (Bergius) Eckl. & Zeyh...... 41 6. Calobota elongata (Thunb.) Boatwr. & B.-E.van Wyk ...... 47 7. Calobota halenbergensis (Merxm. & A.Schreib.) Boatwr. & B.-E.van Wyk...... 50 8. Calobota linearifolia (E.Mey.) Boatwr. & B.-E.van Wyk ...... 54 9. Calobota lotononoides (Schltr.) Boatwr. & B.-E.van Wyk ...... 58 10. Calobota namibensis Boatwr. & B.-E.van Wyk, sp. nov...... 61 11. Calobota obovata (Schinz) Boatwr. & B.-E.van Wyk...... 64 12. Calobota psiloloba (E.Mey.) Boatwr. & B.-E.van Wyk...... 67 13. Calobota pungens (Thunb.) Boatwr. & B.-E.van Wyk...... 70 14. Calobota saharae (Coss. & Durieu) Boatwr. & B.-E.van Wyk ...... 74 15. Calobota sericea (Thunb.) Boatwr. & B.-E.van Wyk...... 77 16. Calobota spinescens (Harv.) Boatwr. & B.-E.van Wyk ...... 82 References ...... 87 Index ...... 93 iv S T R E L I T Z I A 39 (2018)

Abstract

A taxonomic revision of the genus Calobota Thunb., Wiborgia and Wiborgiella. Calo Eckl. & Zeyh. (Fabaceae, Crotalarieae) is pre- bota differs from other closely related genera sented. Sixteen species are recognised in the in the tribe in the following combination of genus. Among these, Calobota namibensis characters: late bark formation, uni- or trifo- Boatwr. & B.-E.van Wyk is described as new liolate to simple, laminar leaves, hairy petals from southwestern Namibia. The anatomy of (C. cuspidosa (Burch.) Boatwr. & B.-E.van the leaves, stems and fruit walls was studied Wyk and C. psiloloba (E.Mey.) Boatwr. & and revealed important characters to distin- B.-E.van Wyk are exceptions), anther configu- guish Calobota from other genera of the tribe ration of 4+5+1 and laterally compressed or Crotalarieae. These include isobilateral leaves terete, usually pubescent pods. A detailed tax- (palisade parenchyma both adaxially and onomic study of the genus is presented here, abaxially) as opposed to dorsiventral leaves including a key to the species, descriptions, (e.g. in Wiborgia Thunb. and Wiborgiella typifications, distributions, phylogenetic rela- Boatwr. & B.-E.van Wyk), and thick-walled tionships and illustrations. fruit with gelatinous fibres in some species. The absence of mucilage cells in the epider- Keywords: Anatomy, Lebeckia, Lebeckia mis of the leaves is also an important distin- sect. Stiza, Fabaceae, Namibia, North Africa, guishing character for the genus with respect Papilionoideae, South Africa, Spartidium, to Aspalathus L., Lebeckia Thunb., Rafnia Wiborgiella. S T R E L I T Z I A 39 (2018) v

Acknowledgements

The authors thank the curators and staff of the listed herbaria for making their specimens available for study or on loan. Financial support from the National Research Foundation of South Africa and the University of Johannesburg are gratefully acknowledged. We are grateful to Drs Sarah K. Gess and Friedrich W. Gess for supplying reprints of their publications on pollinators of the Crotalarieae. Michelle Smith is thanked for making the distribution maps. Collecting permits (permit numbers AAA005-00018-0028, 1362/2009) were provided by the conservation authorities of the Northern and Western Cape, and Namibia.

The authors also wish to thank the anonymous reviewers for providing peer review reports on the manuscript, which resulted in several improvements to the text.

Photographs: Front cover – Calobota angustifolia (typical form) by B.-E. van Wyk Top of back cover – Calobota sericea by J.S. Boatwright Bottom of back cover – Calobota sericea by J.S. Boatwright Inside the cover (collage) – Calobota cytisoides Page vi – Calobota pungens by B.-E van Wyk

New species in this volume: Calobota namibensis Boatwr. & B.-E.van Wyk, sp. nov. (pg. 61)

S T R E L I T Z I A 39 (2018) 1

Introduction

The genus Calobota Eckl. & Zeyh. was re- Aspalathus L., Ezoloba B.-E.van Wyk & Boat- instated to accommodate the 16 species of wr., Rafnia Thunb. and Wiborgia Thunb. the Calobota group (Boatwright et al. 2009), (Boatwright et al. 2008, 2011). Most recently, previously included in the genus Lebeckia the genus Lotononis (DC.) Eckl. & Zeyh. was Thunb. As originally described by Ecklon & also divided into five genera, increasing the to- Zeyher (1836), the genus consisted of two tal number of genera in the Crotalarieae to 16 species, C. cytisoides (Berg.) Eckl. & Zeyh. and (Boatwright et al. 2011). C. pulchella (Andrews) Eckl. & Zeyh., which in actual fact represent the same taxon (now The species of Calobota occur throughout the C. cytisoides). Calobota was later reduced to a Greater Cape Region (Manning & Goldblatt section of an expanded concept of Lebeckia by 2012; Snijman 2013) and extend into Na- Bentham (1844) and several species included mibia. Some are important components of here by Bentham were previously placed in the Sperrgebiet, a species-rich area of south- other genera by Ecklon & Zeyher (1836). western Namibia (Burke & Mannheimer Harvey (1862) followed Bentham’s sectional 2004), while two other species, one described classification in the last revision of the genus herein and C. obovata (Schinz) Boatwr. & Lebeckia where 24 species were recognised. B.-E.van Wyk, are endemic to Namibia (Cra- In Boatwright et al. (2009) Calobota was ven & Vorster 2006). Calobota saharae (Coss. recognised again at genus rank, which also & Dur.) Boatwr. & B.-E.van Wyk, however, included Lebeckia section Stiza E.Mey. and is endemic to North Africa and occurs in the monotypic Spartidium Pomel, based on Algeria, Morocco and Libya (Polhill 1976). a phylogenetic study of the tribe Crotalarieae Calobota can be distinguished from Lebeckia (Boatwright et al. 2008, 2009). The study of and Wiborgiella by a combination of the fol- Boatwright et al. (2008), based on molecular lowing characters: late bark formation, leaves (nrITS and rbcL) and morphological data, in- that are uni- or trifoliolate to simple, laminar; dicated that the genus Lebeckia is not mono- petals hairy (C. cuspidosa (Burch.) Boatwr. phyletic and the division into three genera was & B.-E.van Wyk and C. psiloloba (E.Mey.) subsequently proposed by Boatwright et al. Boatwr. & B.-E.van Wyk are exceptions); an (2009), viz. Lebeckia s.s. with 14 species, Calo anther configuration of 4+5+1; and fruit as bota with 16 species and Wiborgiella Boatwr. laterally compressed or terete, usually pubes- & B.-E.van Wyk with 10 species (Le Roux & cent pods. In contrast, Lebeckia has distinc- Van Wyk 2007, 2008, 2009; Boatwright et tive acicular leaves, glabrous petals and an an- al. 2010; Boatwright & Helme 2014). These ther configuration of 5+5 while Wiborgiella genera form part of the so-called Cape group has brown stems (early bark formation), gla- of genera of the Crotalarieae, along with brous petals and a 4+6 anther arrangement. 2 S T R E L I T Z I A 39 (2018)

Lebeckia has laterally compressed, semi-terete within Calobota are often difficult to distin- or terete pods while the pods are usually in- guish from one another due to the similarity flated in Wiborgiella (Le Roux & Van Wyk between related species and this motivated 2009; Boatwright et al. 2010). The species the need for a modern taxonomic revision. S T R E L I T Z I A 39 (2018) 3

Ethnobotany and chemistry

ETHNOBOTANY Alkaloids Species of Calobota have limited ethnobo- Detailed studies of alkaloids in the Crotalar- tanical uses. Calobota psiloloba has been ieae have made it possible to thoroughly eval- cited as being poisonous, while C. angusti uate their distribution patterns across genera. folia (E.Mey.) Boatwr. & B.-E.van Wyk and Their taxonomic value was discussed by Van C. sericea (Thunb.) Boatwr. & B.-E.van Wyk Wyk et al. (1988) and Van Wyk & Verdoorn are used as remedies for colds (Watt & Breyer- (1990). All genera have been studied except Brandwijk 1962; Van Wyk & Gericke 2000). Bolusia Benth., for which chemical data are Calobota sericea is also popular as firewood in still largely lacking. The chemical structures Namaqualand (Van Wyk & Gericke 2000). of all the various alkaloids are presented in Van Wyk & Verdoorn (1990). The Podalyrie- ae and Crotalarieae differ from all other gen- CHEMISTRY istoids in the absence of a-pyridone alkaloids and from most other tribes in the accumula- The distribution patterns of secondary me- tion of hydroxylated lupanines and their es- tabolites in plants, although sometimes er- ters, especially in their seeds (Van Wyk 2003). ratic, have the potential to provide valuable Within the Crotalarieae, three groups can be insight into phylogenetic relationships. This distinguished based on alkaloid data (Van is especially true for the Fabaceae and more Wyk & Verdoorn 1990; Van Wyk 1991a): specifically the Papilionoideae, where chemi- 1. An unspecialised group without a-pyri- cal patterns are often in agreement with other done alkaloids and without esters of al- evidence (Gomes et al. 1981; Van Wyk 2003; kaloids (Aspalathus, Calobota, Lebeckia, Wink 2003). Some classes of compounds Rafnia, Wiborgia and Wiborgiella, the have been found to be more useful markers so-called Cape group). than others, such as alkaloids and flavonoids in legumes. Several syntheses of the available 2. A group specialising in lupanine-type esters data and literature on alkaloids and flavo- of alkaloids (Euchlora Eckl. & Zeyh., Ezo noids of papilionoid legumes are available loba, Leobordea Del., Listia E.Mey., Pear (e.g. Harborne 1969, 1971; Mears & Ma- sonia Dummer, Robynsiophyton R.Wil bry 1971; Salatino & Gottlieb 1980; Gomes czek and Rothia Pers., i.e. the Pearsonia et al. 1981; Kinghorn & Smolenski 1981; clade). Kinghorn et al. 1982; Hegnauer & Grayer- 3. A group with macrocyclic pyrrolizidine Barkmeijer 1993; Southon 1994; Hegnauer alkaloids (Crotalaria L. and Lotononis s.s. & Hegnauer 1992–2001). (DC.) Eckl. & Zeyh.). 4 S T R E L I T Z I A 39 (2018)

These groups are largely consistent with 2011). The pattern of alkaloids generally sup- those retrieved by phylogenetic studies on ports the new generic classification system for the Crotalarieae based on molecular (nrITS Lotononis (Boatwright et al. 2011). Lotononis and rbcL) and morphological data. The gen- s.s. appears to accumulate only pyrrolizidine era of the Cape group form a monophyletic, alkaloids. The presence of pyrrolizidine alka- although moderately supported clade while loids also supports the sister relationship be- the Pearsonia clade receives strong support tween the Crotalarieae and Genisteae as these (Boatwright et al. 2008), as is also indicated are the only tribes that have genera bearing by the chemical data. Calobota species, much pyrrolizidine alkaloids (Van Wyk 2003). like species of Lebeckia, contain much higher quantities of quinolizidine alkaloids when Flavonoids compared to Wiborgiella species, which contain very low quantities. The latter are more Flavonoids in legumes and within the Cro- similar morphologically to species of Wibor talarieae have not been as well studied as gia in which alkaloids are practically absent alkaloids, possibly due to the somewhat er- (Van Wyk & Verdoorn 1989). ratic patterns found in flavonoid data which limit their utility (Harborne 1971; Van Wyk The presence of macrocyclic pyrrolizidine 2003). The occurrence of isoflavonoids in the alkaloids in Crotalaria and Lotononis s.s. Papilionoideae and the absence of a hydroxyl is unique within the Crotalarieae and was group in position five in all legumes are use- thought to support a sister relationship be- ful characters from flavonoid data (Hegnauer tween these genera (Van Wyk & Verdoorn & Grayer-Barkmeijer 1993; Van Wyk 2003). 1990; Van Wyk 2003). It is now clear, how- Seed flavonoids appear to be more conserva- ever, that the occurrence of these alkaloids tive than those of the leaves (Hegnauer & are rather the result of convergence, as these Grayer-Barkmeijer 1993; De Nysschen et al. genera do not share a sister relationship 1998). (Boatwright et al. 2008). Quinolizidine alkaloids, which are present in most of the other De Nysschen et al. (1998) showed that the genera of the Crotalarieae, are absent from Crotalarieae and the Podalyrieae have the Crotalaria and Lotononis s.s., but present in same four major seed flavonoids (butin, 3’-hy- species of Euchlora, Ezoloba, Leobordea and droxydaidzein, orobol and vicenin-2; chemi- Listia, and they do not co-occur with pyr- cal structures presented in De Nysschen et rolizidine alkaloids in the same species (Van al. 1998), but none of these compounds are Wyk & Verdoorn 1990; Boatwright et al. present in African Genisteae. S T R E L I T Z I A 39 (2018) 5

Phylogenetic relationships

MOLECULAR DATA supported groups were noted. Calobota cus pidosa, C. psiloloba and C. pungens (Thunb.) In Boatwright et al. (2008), 11 species of Calo Boatwr. & B.-E.van Wyk were strongly sup- bota were included in a phylogenetic study ported as being closely related. Calobota cyti of the tribe Crotalarieae. Calobota was weakly soides and C. sericea were strongly supported as supported as monophyletic by separate analy- sister taxa, as were C. angustifolia and C. loton ses of rbcL and ITS, but strongly supported by onoides (Schltr.) Boatwr. & B.-E.van Wyk. The the combined rbcL/ITS and combined rbcL/ expansion of Calobota to include C. saharae ITS/morphological data. Although the sam- (previously the monotypic genus Spartidium) pling was not comprehensive, some strongly was also supported by molecular data. 6 S T R E L I T Z I A 39 (2018)

Horticultural potential

The legume family is the second largest in after scarification or smoke treatment, as is the Cape flora (Manning & Goldblatt 2012) the case with most fynbos legumes (Brown and as such there are several species that & Duncan 2006). Plants of C. cytisoides have become popular horticultural speci- can reach a height of up to 2 m, and with mens. Species of Cyclopia Vent., Indigofera their striking, sweet-scented, yellow flowers L., Liparia L., Podalyria Willd., Psoralea L., should make a bold garden subject. Specific Sutherlandia R.Br. ex Ait. and Virgilia Poir. details on the cultivation of Calobota species are already sold in nurseries, but there are are unavailable, but conditions will likely be several species from other genera, including the same as for other fynbos legumes that are Calobota, which can still be developed as or- presently being cultivated, e.g. Aspalathus, namentals (Brown & Duncan, 2006). Seeds Liparia and Podalyria, which are largely of C. cytisoides can already be purchased at grown in well-drained soil and need water selected seed suppliers and germinate readily during winter in summer-rainfall areas. S T R E L I T Z I A 39 (2018) 7

Materials and methods

Morphological data were obtained from her- a stereoscope (WILD M3Z) with a camera barium specimens as well as fresh plant ma- lucida attachment, and drawings were done terial from field excursions. Specimens were by the first author. preserved in FAA (Sass 1958; formaldehyde: acetic acid: 96% alcohol: water; 10:5:50:35) Data on the mapped distribution of the vari- and silica (Chase & Hill 1991). All these col- ous species were gathered from the herbari- lections are housed in the University of Jo- um material, field notes and from Leistner hannesburg Herbarium (JRAU). Specimens & Morris (1976, for southern African taxa). of Calobota from the following herbaria were The data for each species, with the exception also studied: BM, BOL, GRA, J, JRAU, K, of C. saharae, are recorded using the Quar- NBG (including SAM and STE), P, PRE, S, ter Degree Reference System (Edwards & SBT, UPS and WIND. Online photographs Leistner 1971; also Leistner & Morris 1976). of the collections of B, M, TCD and Z were The basic unit used in this system is the one- degree square of latitude and longitude, also studied. The material was arranged ac- which is designated by a Degree Reference cording to geographical distribution so that Number (degrees of latitude and longitude the relevant measurements and observations of the northwest corner) and the district could be done within several geographical name of that square. populations. At least three measurements per specimen were done where possible. Mature For anatomical studies, fresh material was flowers from material fixed in FAA or flowers fixed in FAA or dried material was rehydrated rehydrated from rich herbarium specimens by briefly boiling in distilled water and then were dissected and mounted in glycerol. Spec- fixing in FAA for 24 hours. The method of imens from which dissections were made are Feder & O’Brien (1968) for embedding in listed in Boatwright (2009). Floral characters glycol methacrylate (GMA) was used, except recorded were: pedicel length, length of the that the final infiltration in GMA was done bracts and bracteoles, flower length, length of for a minimum of five days. Sections were the calyx, length of the calyx tube, depth of stained by the periodic acid Schiff/toluidine the upper, lateral and lower sinuses of the ca- blue (PAS/TB) staining method (Feder & lyx, width of the upper, lower and lateral ca- O’Brien 1968) and mounted. Voucher in- lyx lobes, length and width of the standard-, formation for the material used in the ana- wing- and keel petals and the length of the tomical studies can be found in Boatwright claws of each petal. Drawings of vegetative (2009). Micrographs were taken with a JVC and reproductive structures were done using KY-F1030 digital camera. 8 S T R E L I T Z I A 39 (2018)

Vegetative morphology and anatomy

HABIT AND BRANCHES deposits is initiated early, giving even young stems a brownish colour (Boatwright et al. All species of Calobota are perennial, multi- 2010). Some species of Calobota, such as stemmed shrubs usually about 1.0–1.5 m, C. pungens and C. saharae, lack leaves on but C. cuspidosa can reach a height of up to mature branches and in these it is likely that 4 m (Figure 1). Calobota lotononoides is a de- the stem performs the main photosynthetic cumbent shrublet, to 0.5 m in height, and has function. The epidermis in most species is extensive underground branches. This spe- covered by a thick cuticle and the epidermal cies appears to be adapted to sandy habitats cells are bottle-shaped (Figure 2). as the underground branches allow the stems and leaves to emerge above ground when they become buried (Boatwright & Van Wyk LEAVES 2007). Calobota acanthoclada (Dinter) Boat- wr. & B.-E.van Wyk, C. cuspidosa, C. psiloloba, Leaves in Calobota vary from trifoliolate or C. pungens and C. spinescens (Harv.) Boatwr. rarely five-foliolate to unifoliolate or sim- & B.-E.van Wyk are spinescent shrubs, with ple. In three species with unifoliolate leaves, the branch ends and inflorescences terminat- C. cuspidosa, C. psiloloba and C. pungens, ing in spines. The typical form of C. angusti the juvenile leaves are trifoliolate and be- folia sometimes has spinescent branches, but come unifoliolate on the mature branches is more often not spinescent. (Dahlgren 1970; Polhill 1976; Boatwright et al. 2009). A similar reduction has been ob- The young branches in species of Calobota, served in C. obovata, which also has unifoliol- with the exception of C. acanthoclada, are ate leaves restricted to the mature branches. green as opposed to the brown colour found Whether those of species with simple leaves in most species of Wiborgiella (Boatwright have arisen in the same way, as suggested by et al. 2010). This is due to the presence of Dahlgren (1970), should be investigated by several chlorenchyma layers below the epi- studying the juvenile leaves of these species. dermis (Figure 2), a feature usually found The leaves of Calobota species are invariably in papilionoid legumes when the leaflets are laminar, rather than acicular and phyllodi- reduced or absent (Metcalfe & Chalk 1950). nous as found in Lebeckia, usually with long, Although green stems are also found in other slender petioles. Blade margins in Calobota genera of the Crotalarieae, notably Crota are entire and stipules are lacking. The leaves laria, Lebeckia, Lotononis and Rafnia, it is are always pubescent or at least pilose. The a useful character to distinguish Calobota leaf arrangement in Calobota is alternate and from Wiborgiella and Wiborgia. In these last widely spaced, but in C. acanthoclada and two genera, a periderm with tanniniferous C. spinescens the leaves are fasciculate. This S T R E L I T Z I A 39 (2018) 9

A B

C D

Figure 1.—Habit of Calobota species. A, C. cinerea; B, C. cytisoides; C, C. halenbergensis; D, C. lotononoides; [on next page]: E, C. pungens; F, C. cuspidosa; G, C. spinescens; H, C. sericea. Photographs: A–E, G–H: J.S. Boatwright; F: B.-E. van Wyk. 10 S T R E L I T Z I A 39 (2018)

E F

G H

Figure 1. (cont.)—Habit of Calobota species. [previous page]: A, C. cinerea; B, C. cytisoides; C, C. halenbergen sis; D, C. lotononoides; [this page]: E, C. pungens; F, C. cuspidosa; G, C. spinescens; H, C. sericea. Photo- graphs: A–E, G–H: J.S. Boatwright; F: B.-E. van Wyk. S T R E L I T Z I A 39 (2018) 11

A B

C D

Figure 2.—Transverse sections through the stems and leaves of Calobota species. A, portion of the stem of C. spinescens showing several layers of chlorenchyma below the epidermis (Boatwright et al. 158, JRAU); B, portion of the stem of C. angustifolia showing several layers of chlorenchyma below the epidermis (Boatwright et al. 138, JRAU); C, portion of the juvenile leaf of C. pungens showing the presence of mucilage cells in the epidermis (Boatwright et al. 106, JRAU), D, portion of the leaf of C. obovata showing the isobilateral mesophyll (Kers 152, WIND). Scale bars: A–C, 0.1 mm; B, 0.2 mm.

is a useful character, especially to distinguish Anatomical study of the leaves of Calobota C. spinescens from C. angustifolia. Leaflet shape species (Figure 2) provided contrasting char- varies from oblanceolate to elliptic or obovate acters with those of Lebeckia and Wibor to suborbicular and less often spathulate. The giella. In Calobota, the leaves are isobilat- apices may be acute, obtuse, retuse, trun- eral and lack mucilage cells in the epidermis. cate, mucronulate or recurved-mucronate, Transverse sections through juvenile leaves of and the base of the leaflets angustate, attenuate C. pungens show mucilage cells in the epider- or cuneate. Calobota angustifolia is a highly mis (Figure 2), suggesting that there is a devel- variable species and four regional forms can be opmental loss of these cells in Calobota. Calo distinguished, mainly on leaf size, shape and bota saharae is an exception in that mucilage especially the width of the leaflets. cells remain present in mature leaves. Gregory 12 S T R E L I T Z I A 39 (2018)

& Baas (1989) speculate that these cells are show the palisade parenchyma adjacent to usually associated with plants that occur in the epidermis and mucilage cells are present. Mediterranean climates, and consistent with Wiborgiella species have dorsiventral leaves this, C. saharae is endemic to Mediterranean (with both palisade and spongy parenchyma) North Africa. In Lebeckia the acicular leaves and mucilage cells are present in the epidermis. S T R E L I T Z I A 39 (2018) 13

Reproductive morphology and anatomy

INFLORESCENCE Bracts and bracteoles vary little among species. A single bract is located at the base of the The inflorescence in Calobota is fairly uni- pedicel, with two bracteoles further up near form in structure, and all species have termi- the middle of the pedicel. These bracts and nal racemes that vary only in length and in the bracteoles are small, linear and adaxially pu- number of flowers per raceme (Figure 3). Ter- bescent, except in C. cytisoides and C. obovata minal racemes are rather common in Papilio- where they are often ovate. noideae and also in the Crotalarieae (Polhill 1976). Notably in C. elongata (Thunb.) In the flower, the calyces of Calobota species Boatwr. & B.-E.van Wyk, the inflorescence is are typically of the lebeckioid type (Polhill a densely flowered spike, with more than 100 1976) and are subequally lobed, with the up- sessile flowers (Dahlgren 1967). per sinus often deeper than the lateral or lower sinuses and the carinal lobe consistently narrower than the others. The outer surface FLOWERS of the calyx is usually pubescent, pilose or at Flower size is somewhat diagnostic at spe- least glabrescent. The tips of the calyx lobes cies level in Calobota. The largest flowers are minutely pubescent on the inner surface, are found in C. cytisoides and C. sericea (to which is found in most of the genera of the 35 mm and 24 mm long, respectively) and Cape group of the Crotalarieae, with the no- C. cuspidosa has flowers to 20 mm long. Re- table exception of Aspalathus. maining species range from 17 mm long in C. linearifolia (E.Mey.) Boatwr. & B.-E.van The standard petal is mostly widely ovate Wyk and C. saharae to less than 12 mm long in the species of Calobota, but is elliptic in in C. elongata, C. halenbergensis (Merxm. C. elongata, orbicular in C. halenbergensis & A.Schreib.) Boatwr. & B.-E.van Wyk, and transversely oblong in C. lotononoides. C. namibensis, C. psiloloba and C. angustifo The apices are either acute or obtuse, except lia. The consistently yellow petals (Figure 3) in C. psiloloba and C. lotononoides, which are usually pubescent in Calobota or at least have emarginate apices. The claws are rela- pilose along the dorsal midrib of the standard tively short and less than 6 mm long, except in petal, with the exception of C. cuspidosa and C. cuspidosa, C. cytisoides and C. lotononoides C. psiloloba that have totally glabrous petals. where the claws are longer. Basal callosi- Floral pedicels are short in most species, but ties are only found only in C. cytisoides. The may be longer in C. cytisoides (to 11 mm) standard petal is usually pubescent in Calo and C. psiloloba (to 6 mm) or absent in bota species or at least with pilose hairs along C. elongata. the dorsal midrib, except in C. cuspidosa 14 S T R E L I T Z I A 39 (2018)

Figure 3.— Flowers and fruit of Calobota species. A, C. cytisoides; B, C. sericea; C, C. pungens; D, C. cinerea; E, C. cuspidosa; F, C. angustifolia. Photographs: A, C–E: B.- E. van Wyk; B, F: J.S. Boatwright. S T R E L I T Z I A 39 (2018) 15

C D

E F 16 S T R E L I T Z I A 39 (2018) and C. psiloloba, which have totally glabrous The value of staminal characters in the gen- standard petals. istoid legumes is well known (Polhill 1976; Schutte & Van Wyk 1998; Boatwright et al. The shape of the wing petals is fairly uniform 2009) and provide reliable characters to segre- in the genus and mostly oblong with obtuse gate Calobota and Wiborgiella from Lebeckia apices. They are either shorter than the keel s.s (Boatwright et al. 2009). The species of petals (Calobota cinerea (E.Mey.) Boatwr. & Calobota have a 4+5+1 anther arrangement B.-E.van Wyk, C. cuspidosa, C. linearifolia, (four long basifixed anthers, five short dorsi- C. namibensis, C. obovata, C. psiloloba, C. pun fixed anthers and an intermediate carinal an- gens), as long as or shorter than the keel petals ther), while those of Lebeckia have a 5+5 ar- (C. acanthoclada, C. cytisoides, C. spinescens, rangement (the carinal anther resembles the C. sericea), as long as or longer than the keel long basifixed anthers) and species of Wibor petals (C. elongata, C. saharae, C. angustifo giella have a 4+6 arrangement (the carinal an- lia) or markedly longer than the keel petals ther resembles the short dorsifixed anthers). (C. halenbergensis, C. lotononoides). Calobota lotononoides is unique within the genus in The pistil in Calobota is subsessile or shortly having the lamina of the wing petals twice as stipitate, usually with a linear ovary and up- long as those of the keel petals; the wing petal curved style. The ovary may be either pubes- laminas of the other species are never more cent or glabrous, while the style is always gla- than 1.3 times longer than those of the keel brous. The ovary may contain as few as three petals (Boatwright & Van Wyk 2007). The to six ovules (C. elongata, C. namibensis) or as wing petals are glabrous in most species, but many as 34 ovules (C. cytisoides). The style is pilose in C. acanthoclada, C. cinerea, C. line longer than the ovary in C. namibensis. arifolia, C. obovata and C. pungens. Sculptur- ing is present in all species on the upper basal The nectar of Calobota species is generally and upper central parts of the wing petals. In sucrose-rich (Van Wyk 1993), typical of most some species, e.g. C. cytisoides, C. elongata and bee-pollinated legumes. The potential polli- C. sericea, the sculpturing extends to the low- nators of some of the Cape genera of the Cro- er basal, lower central and upper/lower distal talarieae were studied by Gess & Gess (1994). parts of the wing petals (cf. Stirton 1981). They found that the flowers of C. spinescens and C. sericea were visited by solitary aculeate Keel petals are boat-shaped with obtuse apices Hymenoptera, specifically Masarinae and and pockets are invariably present in all Calo Megachilinae of the tribes Megachilini and bota species. The petals are either pubescent Anthidiini. Some species of Xylocopinae also (C. acanthoclada, C. cinerea, C. linearifolia, occasionally visited C. sericea and Apis mel C. cytisoides, C. namibensis, C. obovata, C. pun lifera L. was also often abundant. The meg- gens and C. sericea) or glabrous (C. psiloloba, achiline and masarine visitors were able to C. cuspidosa, C. elongata, C. halenbergensis, trip (displacement of the keel petals to come C. lotononoides, C. saharae, C. angustifolia). In into contact with the pollen) the flowers of C. spinescens the keel petals may be either pilose C. spinescens, but not those of C. sericea sug- or glabrous. gesting that they are potential pollinators of S T R E L I T Z I A 39 (2018) 17 only the former. Apis mellifera, however, was FRUIT the only species that was able to trip the flowers of Calobota sericea making it a potential The pods seen in Calobota species vary from pollinator of this species (Gess & Gess 1994, laterally compressed and membraneous to 2004, 2006). The diversity of flower visits to terete and glabrous or pubescent. The pods other species of Calobota and the Crotalarie- are generally many-seeded, with the excep- ae has been recorded by Gess & Gess (2003). tion of C. elongata where the pods are small,

A B

C D

Figure 4.—Transverse sections through the fruit walls of Calobota species showing thick-walled (A, C) and thin-walled (B, D) pods. Note the mass of multicellular hairs associated with the endocarp in A, C. A, C. angustifolia (Boatwright et al. 138, JRAU); B, C. spinescens (Boatwright et al. 158, JRAU); C, C. sericea (Boatwright et al. 151, JRAU); D, C. elongata (Van Wyk 2562b, JRAU). Scale bars: A–D, 0.2 mm 18 S T R E L I T Z I A 39 (2018) light and usually single-seeded. In this spe- protective function. In the species with thin- cies, the petals dry out but are persistent and walled pods, the mesocarp is thinner and has serve as wings for the disperal of the tiny pod only one or two rows of sclerenchyma cells by wind, through a rolling action in the open adjacent to the endocarp (Figure 4). One or sandy places where these plants typically rarely two layers of cells make up the endo- grow. A similar adaptation to wind dispersal carp and the sponge-like dense mass of hairs is found in Leobordea sect. Synclistus (B.-E. is absent. The segregation of Lebeckia s.s. into van Wyk) B-E.van Wyk & Boatwr. (Van Wyk three genera is supported by fruit anatomical 1991b). data (Le Roux et al. 2011).

Anatomically there are two types of fruit walls in species of Calobota, distinguished by SEEDS their thickness (Figure 4) (type I and III according to Le Roux et al. 2011). It is also one In Calobota the seeds are reniform or oblong- of only two genera in Crotalarieae in which reniform and vary from light brown to pink, gelatinous fibres are found in the endocarp, sometimes with brown mottling, and gen- which is associated with a dense mass of mul- erally have a brown hilum. The surface is ticellular hairs which give the appearance of smooth in all but one species, namely C. loto a white sponge to the naked eye and seem nonoides, which has rugose seeds (Boatwright to envelop the seed, perhaps to serve some & Van Wyk 2007). S T R E L I T Z I A 39 (2018) 19

Taxonomic treatment

Calobota Eckl. & Zeyh., Enum. Pl. Afric. Austral. [Ecklon & Zeyher] 2: 192. [Jan.] 1836; Boat- wr. & B.-E.van Wyk, S. African J. Bot. 75: 553. 2009. Lebeckia sect. Calobota (Eckl. & Zeyh.) Benth., London J. Bot. 3: 358–361. 1844. p.p. LECTOTYPE (designated by Boatwright et al. 2009: 552): Calobota cytisoides (Bergius) Boatwr. & B.-E.van Wyk. Acanthobotrya Eckl. & Zeyh., Enum. Pl. Afric. Austral. [Ecklon & Zeyher]. 2: 192. [Jan] 1836, p.p. LECTOTYPE (designated by Boatwright et al. 2009: 552): A. pungens Eckl. & Zeyh. [= Calobota psiloloba (E.Mey.) Boatwr. & B.-E.van Wyk]. Stiza E.Mey., Comm. Pl. Afr. Austral. [Meyer] 1: 31. [Feb.] 1836. Lebeckia sect. Stiza (E.Mey.) Benth., London J. Bot 3: 355, 1844. LECTOTYPE (designated by Boat- wright et al. 2009: 552): Stiza erioloba E.Mey. [= Calobota pungens (Thunb.) Boatwr. & B.-E. van Wyk]. Spartidium Pomel, Nouv. Mat. Fl. Atl.: 173. 1874. TYPE: S. saharae (Coss. & Durieu) Pomel.

[Note: As mentioned by Bentham (1844), Acanthobotrya is a mixed concept representing at least four different genera, including species of Calobota, Wiborgia and Wiborgiella. However, the diag- nosis agrees with the concept of Stiza E.Mey. in the linear-oblong, compressed fruits.].

Woody, sometimes spinescent shrubs or to obovate, pubescent, caducous; bracteoles shrublets; branches thick and woody; young linear, pubescent, caducous. Flower with co- branches green, with chlorenchyma and lack- rolla yellow, usually pubescent or at least pi- ing bark (except in C. acanthoclada), pubes- lose along the median section of the standard cent, often sericeous. Stipules absent. Leaves petal (except in C. cuspidosa and C. psiloloba); unifoliolate or digitately trifoliolate (rarely calyx subequally lobed, upper sinus often 5-foliolate), sometimes caducous so that ma- deeper than the lateral or lower sinuses, cari- ture plants appear leafless, seasonally decidu- nal lobe narrower than the others, pubescent ous in some species; petioles shorter or longer or at least glabrescent, tips of lobes minutely than leaflets, pubescent; leaflets oblanceolate pubescent on inner surface; standard lin- to elliptic or obovate to suborbicular, less of- ear to widely ovate, with basal callosities in ten spathulate, pubescent. Inflorescence ter- C. cytisoides; wing petals narrowly oblong minal, few to multi-flowered racemes or rare- to oblong or slightly ovate, longer or shortly a spike; pedicels pubescent; bracts linear er than the keel, terminal parts sometimes 20 S T R E L I T Z I A 39 (2018) pubescent; apex obtuse; keel petals oblong, bark formation is early in most species of pockets invariably present, terminal parts Wiborgiella, so that even the young twigs sometimes pubescent; apex obtuse; anthers are not green but covered in brown bark (this dimorphic, four long, basifixed anthers al- is also true of Wiborgia species). The green ternating with five ovate, dorsifixed anthers, twigs are a useful diagnostic character, vis- carinal anther intermediate in size and shape ible even in sterile herbarium specimens. between the basifixed and dorsifixed anthers; Calobota also differs from Wiborgiella in the pistil subsessile to very shortly stipitate; ovary hairy petals (C. cupidosa and C. psiloloba are linear to slightly elliptic, with four to many exceptions) seeing that the petals are glabrous ovules, pubescent or glabrous; style longer or in Wiborgiella. The most reliable diagnos- shorter than the ovary, curved upwards, gla- tic character to distinguish Calobota from brous. Fruits as pods laterally compressed or Wiborgiella is the anther configuration of terete, linear to oblong, few- to many-seeded, 4+1+5 (4+6 in the latter). The pods are semi- glabrous or pubescent, dehiscent or inde- terete or laterally compressed in Calobota and hiscent; endocarp glabrous or hairy. Seeds usually pubescent, whereas pods are terete reniform to oblong-reniform, or less often or rarely laterally compressed in species of suborbicular; colour variable, light pink to Wiborgiella and always glabrous. pink, sometimes mottled with brown; hilum round, brown or black; surface smooth (ru- Notes on distribution gose only in C. lotononoides; seeds of C. na mibensis, C. obovata and C. saharae not seen). The species of Calobota occur in the Eastern, Northern and Western Cape provinces, with Diagnostic characters some restricted to Namibia. One species, C. saharae, is endemic to North Africa where Bark formation is late, so that the twigs re- it occurs on sand dunes from Libya to Algeria main green and photosynthesising, while and Morocco (Polhill 1976). S T R E L I T Z I A 39 (2018) 21

KEY TO THE SPECIES OF CALOBOTA

1a. Plants strongly spinescent shrubs; pods laterally compressed, often falcate ...... 2 2a. Plants leafless shrubs, if leaves present then widely separated ...... 3 3a. Petals pilose; ovary pubescent; pods densely tomentose ...... 13. C. pungens 3b. Petals glabrous; ovary glabrous; pods glabrous ...... 4 4a. Flowers usually (15–)16–18(–20) mm long; pods (21–)28–49(–53) mm long; plants restricted to the Northern Cape ...... 4. C. cuspidosa 4b. Flowers usually 8–11 mm long; pods 14–17(–33) mm long; plants restricted to the Eastern Cape ...... 12. C. psiloloba 2b. Plants shrubs with leaves present, leaves fasciculate ...... 5 5a. Leaves trifoliolate; young stems green; pods constricted between the seeds ...... 16. C. spinescens 5b. Leaves simple; young stems brown; pods not constricted between the seeds ...... 1. C. acanthoclada 1b. Plants not spinescent (or if rarely spinescent then pods terete); pods terete, if rarely laterally compressed then plants not spinescent, but erect or virgate, leafless shrubs restricted to North Africa ...... 6 6a. Leaflets widely obovate or orbicular, 8–25 mm wide ...... 7 7a. Leaves trifoliolate; inflorescences long (80–432 mm), spicate and densely flowered (more than 100 flowers); pods ovate to elliptic, 5–8 × 1.5–4.0 mm, 1- to 2-seeded ...... 6. C. elongata 7b. Leaves unifoliolate (basal leaves rarely trifoliolate); inflorescences short (24–85 mm), racemose and 7- to 16-flowered; pods linear, 30–40 × 4–5 mm, 5- to 7-seeded ...... 11. C. obovata 6b. Leaflets elliptic to oblanceolate, linear, spathulate, or rarely ovate or obovate, less than 10 mm wide (if leaflets elliptic to oblanceolate and rarely more than 10 mm wide then plants large shrubs up to 2 m with large flowers, 18–35 mm long) ...... 8 8a. Wing petals longer than keel ...... 9 9a. Plants pilose and not sericeous; pods glabrous; pistil glabrous ...... 7. C. halenbergensis 9b. Plants sericeous; pods pubescent or sericeous; pistil pubescent or sericeous ...... 10 10a. Leaflets not folded inward and directed to the same side; flowers 7–11 mm long; wing petals up to 1.3 × as long as the keel ...... 2. C. angustifolia 10b. Leaflets conduplicate and secund; flowers 12–15 mm long; wing petals 2 × as long as the keel ...... 9. C. lotononoides 8b. Wing petals shorter than keel ...... 11 11a. Leaves trifoliolate and petiolate ...... 12 12a. Flowers medium-sized (11–16 mm long); standard petal reflexed; wing petals with sculpturing restricted to the upper basal and upper central parts; pods tomentose ...... 3. C. cinerea 12b. Flowers generally large (between 13 and 35 mm long); standard petal upright; wing petals with sculpturing extending to the lower basal, lower central and upper or lower distal parts; pods glabrous or sericeous ...... 13 13a. Leaflet apices mucronulate; calyx glabrous; ovary glabrous; pods glabrous ...... 5. C. cytisoides 13b. Leaflet apices acute; calyx sericeous; ovary sericeous; pods sericeous ...... 15. C. sericea 11b. Leaves simple and sessile ...... 14 14a. Leaflets and stems sericeous or silky; pods short-sericeous ...... 8. C. linearifolia 14b. Leaflets and stems not sericeous; pods glabrous or densely pubescent ...... 15 15a. Leaflets elliptic to ovate; keel petals glabrous; pistil glabrous; pods glabrous; restricted to North Africa ...... 14. C. saharae 15b. Leaflets linear; keel petals densely pubescent; pistil densely pubescent; pods densely pubescent; restricted to Namibia ...... 10. C. namibensis 22 S T R E L I T Z I A 39 (2018)

1. Calobota acanthoclada (Dinter) Boatwr. & B.-E.van Wyk, S. African J. Bot. 75: 553. 2009. Basionym: Lebeckia acanthoclada Dinter, Repert. Spec. Nov. Regni Veg. 30: 196. 1932. TYPE: NAMIBIA, Granietberge [2715 AD], Dinter 6694 (lectotype, designated here, B, photo!). Lebeckia spathulifolia Dinter, Repert. Spec. Nov. Regni Veg. 30: 197. 1932. TYPE: NA- MIBIA, Kleinfonteiner Fläche [2516 BB], Dinter 3735 (holotype, B, photo!; isotypes, BM!, PRE!, SAM!, Z, photo! 2 sheets). Lebeckia candicans Dinter, Repert. Spec. Nov. Regni Veg. 30: 198. 1932. TYPE: NAMIB- IA, Kleinfonteiner Fläche [2516 BB], Dinter 3737 (holotype, B, photo!; isotypes, BM!, BOL!, NBG!, PRE!, S!, Z, photo!).

Small, erect or decumbent, multi-stemmed, style shorter than ovary, 3.5–5.5 mm long. spinescent shrub up to 1 m in height. Branches Pods linear, laterally compressed, sericeous, all brown; young branches slightly sericeous to subsessile to shortly stipitate, 12–19 × 1.5– pilose, not green; older branches covered with 3.0 mm, ±4- to 5-seeded, dehiscent; endocarp brown bark, pilose. Leaves simple, persistent; glabrous. Seeds oblong-reniform to suborbicu- petiole absent; fasciculate, sericeous on both lar, 1.5–2.5 × 1.0–2.5 mm, mature seeds light surfaces, spathulate, sessile, 6–20 × 1–2 mm, brown to orange, surface smooth (Figure 5). apex recurved-mucronate, base angustate. In Flowering time: flowering specimens have been florescence 12–25 mm long, racemose, with collected from February through to October. 3 to 5 flowers; pedicel 2–3 mm long; bract 1.5–2.5 mm long, linear, sericeous; bracteoles 0.5–1.0 mm long, linear, sericeous. Flowers Distribution and habitat 9–12 mm long, bright yellow. Calyx 5–7 mm Calobota acanthoclada is mainly found in Na- long, densely sericeous; tube 2.5–4.0 mm long; mibia but extends into the Richtersveld of lobes 2–4 mm long, subulate. Standard 9.5– South Africa (Figure 6). A collection that was 14.0 mm long; claw linear, 2–5 mm long; lami- made in the Cederberg (Western Cape Prov- na ovate, 6.5–8.5 × 7–8 mm; apex acute; dorsal surface densely sericeous. Wings 9.5–11.0 mm ince) suggests that the species extends even fur- long; claw 2.5–3.5 mm long; lamina oblong, ther south into South Africa. It grows on sandy as long as or shorter than keel, 6.0–7.5 × 2.5– loam, rocky soil, limestone or dolomitic lime- 3.0 mm, pilose, with 5–7 rows of sculpturing. stone at altitudes of between 170 and 1 300 m. Keel 10.0–12.5 mm long; claw 3–4 mm long; lamina boat-shaped, 6.5–8.5 × 3.5–4.5 mm, IUCN Red List category pilose to sericeous on terminal parts. Pistil subsessile to shortly stipitate, pubescent; ovary lin- Considered Endangered (EN) by Raimondo ear, 7.0–9.5 × 0.7–1.0 mm with 9 to 16 ovules; et al. (2009). S T R E L I T Z I A 39 (2018) 23

A B C

D E F

G1 G2 J1 J2

I1 I2 I3 K2

Figure 5.—Morphology of Calobota acanthoclada. A, flower in lateral view; B, standard petal; C, outer surface of the calyx (upper lobes to the left); D, wing petal; E, keel petal; F, pistil; G1, bract; G2, bracteoles; H, androecium; I1, long, basifixed anther; I2, intermediate carinal anther; I3, short, dorsifixed anther; J1, J2, leaves in abaxial view; K1, pod in lateral view; K2, pod in dorsal view. A–G: Mannheimer CM2012a, WIND; H–I: Williamson 3376, BOL; J: Mannheimer et al. CM 899, WIND; K: Pillans 5149, BOL. Scale bars: A–I, 1 mm; J, K, 10 mm. 24 S T R E L I T Z I A 39 (2018)

Additional specimens examined NAMIBIA. 2516 (Helmeringhausen): Kleinfonteiner Flache (–BB), Dinter 6269 (B, BOL). 2518 (Tses): Kovisberge (–BB), Dinter 6293 (B, BM); Merxmül ler & Giess 28445 (PRE, WIND). 2615 (Lüderitz): Klinghardt Mountains (–CA), Clark & Müller 345 (WIND); 1.0 ml. [1.61 km] south of Lagoon, Lüderitz Bay (–CA), Giess & Van Vuuren 669 (K, PRE, WIND); between Elizabeth Bay and Kolmanskop, towards coast along old mine road (–CA), Mannheimer CM2012a (WIND); Lüderitz (–CA), Merxmüller & Giess 3076 (PRE, WIND); Lüderitz, on road to Diazpunt (–CA), Müller & Jankowitz 270 (WIND); proposed wind Figure 6.—Distribution of Calobota acanthoclada. farm site, east of Grosse Bucht (–CC), Mannheimer & Burke CM 1700 (WIND); Grillenthal, blue dolomite hills on the road south to Oranjemund from Lüderitz in the Diamond Area No. 1 (–CD), Bean & Oliver 2425 (BOL, NBG); Sperrgebiet South, 1.5 km north of Discussion Grillenthal (–CD), Jürgens 28103 (PRE). 2616 (Aus): 56 mls [90.1 km] west of Aus, Schotterhang (–CB), This species is similar to C. spinescens in Leippert 4080 (WIND). 2715 (Bogenfels): ca. 5 km the fasciculate leaves, ovate and sericeous north of Pomona (–AB), Burgoyne 8270 (PRE); Pomo- standard petal and spinescent habit, but na (–AB), Dinter 6366 (BM, BOL, NBG 2 sheets, K, the branches are brown and not green as in PRE, S, WIND); along road to Pomona (–AB), Mann C. spinescens (and indeed all other species of heimer et al. CM899 (WIND); Rote Kuppe, Chamais road (–BC), Bartsch SB 1043 (WIND); 13 km west Calobota). It also differs in the simple leaves of Bogenfels houses, at ridge running south of road (trifoliolate in C. spinescens), smaller flowers (–BC), Mannheimer CM961 (WIND); Klinghardt and shorter calyx lobes. The pods are not Mountains (–BD), Dinter 3980 (BOL, PRE); Boegoe- constricted between the seeds as is the case in berg (–DD), Dinter 6574 (B, photo!, BM!); Mann C. spinescens. heimer CM1015 (WIND); summit of the Boegoeberg (–DD), Williamson 2594a (BOL 2 sheets). Dinter 6694 cited by Dinter (1932) is still SOUTH AFRICA. 2816 (Oranjemund): Brandkaros extant in B and is here chosen as lectotype. (–BC), Venter 8863 (PRE); 0.5 km east of Brandkaros Dinter (l.c.) cited only a single collection on road to Anniesfontein (–DA), Botha 3201 (PRE); (Dinter 3735) in the original description of Richtersveld, Kortdoorn (–DA), Jürgens 22585 (PRE); Witbank (–DC), Pillans 5149 (BOL, K). 3219 (Wup- Lebeckia spathulifolia and also only one col- pertal): Oudekraal farm between Ramkraal and Ou lection (Dinter 3737) in the description of kraal, Cederberg (–AD), Koekemoer 2467 (PRE). Pre- Lebeckia candicans. The specimens in B of cise locality unknown: Nautilus, Kinges 2580 (PRE); these collections are still extant. Summit of Kortderm, Williamson 3376 (BOL). S T R E L I T Z I A 39 (2018) 25

2. Calobota angustifolia (E.Mey.) Boatwr. & B.-E.van Wyk, S. African J. Bot. 75: 553. 2009. Basionym: Lebeckia angustifolia E.Mey., Linnaea 7: 155. 1832. Acanthobotrya angustifolia (E.Mey.) Eckl. & Zeyh., Enum. Pl. Afric. Austral. [Ecklon & Zeyher] 2: 194. Jan. 1836. TYPE: SOUTH AFRICA, s. loc., Ecklon s.n. (lectotype, designated here, S!). Spartium sericeum Ait. in Hortus Kew. [W.Aiton] 3: 12. 1789, non Lebeckia sericea Thunb. TYPE: SOUTH AFRICA, Cape of Good Hope, Masson s.n. sub BM000794145 (lectotype, designated here, BM, photo!, specimen on the left). Lebeckia multiflora E.Mey., Comm. Pl. Afr. Austral. [Meyer] 1: 34. Feb. 1836. TYPE: SOUTH AFRICA, betw. Holrivier & Mierenkasteel [3118 CB], Drège 6474 ‘III, C’ (lectotype, designated here, P!; isolectotypes, BM!, K!, PRE!, S!). Lebeckia decutiens E.Mey. var. a canescens E.Mey., Comm. Pl. Afr. Austral. [Meyer] 1: 34. Feb. 1836. TYPE: SOUTH AFRICA, near Heerenlogement [3118 DC], Drège s.n. ‘III, E, a’ (lectotype, designated here, P!; isolectotype, BM!, K!). Lebeckia multiflora var. parviflora Schinz in Abhandl. Bot. Ver. Prov. Brandenb. 30: 157. 1888. Lebeckia parvifolia (Schinz) Harms in Engl. Pflanzenw. Afr. 3(1) (Engl. & Drude, Veg. der Erde 9): 545 (1915). TYPE: NAMIBIA, Angra Pequena [2615 CA], Schinz 811 (lectotype, designated here, Z, photo!; isolectotype, BOL!, K!).

Erect or spreading, multi-stemmed, unarmed linear, 2.0–4.5 mm long; lamina widely ovate, or rarely spinescent shrub up to 2 m in height. 4–9 × (3.8–)4.5–10.0 mm, pilose along dorsal Branches green; young branches sericeous, midrib; apex obtuse. Wings 7–13(–14) mm older branches sericeous or glabrous with long; claw 2.5–4.5(–5.5) mm long; lamina brown bark. Leaves digitately trifoliolate, very oblong, as long as but more often longer than variable, deciduous; petiole longer or shorter keel, 4.5–8.5 × 2.0–4.5 mm, glabrous, with than leaflets, 8–45 mm long; leaflets linear 4–12 rows of sculpturing. Keel 6.5–12.5 mm to oblanceolate, alternate, sericeous, subses- long; claw 2.5–4.5(–5.5) mm long; lamina sile, terminal leaflet 5–30 × 0.5–5.0 mm, boat-shaped, 4–8 × 2.5–5.0 mm, glabrous. lateral leaflets 4–25 × 0.5–4.0 mm, apex Pistil subsessile to shortly stipitate, pubes- acute, base cuneate. Inflorescence 20–200 mm cent; ovary linear, 5.0–9.5 × 0.5–1.1 mm long, with 7 to 40 flowers; pedicel 1–2 mm with 11 to 20 ovules; style shorter than long; bract 1–3 mm long, linear, pubescent; ovary, 2.5–5.0 mm long. Pods linear, terete, bracteoles 0.5–2.0 mm long, linear, pubes- sericeous, subsessile to shortly stipitate, 18– cent. Flowers 7–11 mm long, bright yellow. 35 × 2–4 mm, 5- to 12-seeded, dehiscent; Calyx 4.0–6.5 mm long, pubescent; tube endocarp hairy. Seeds oblong-reniform, 2.5– 2.5–4.5 mm long; lobes 1.0–3.5 mm long, 3.5 × 1.5–2.0 mm, mature seeds light brown deltoid. Standard 6.5–13.0 mm long; claw or light orange, sometimes mottled with 26 S T R E L I T Z I A 39 (2018)

A1 A2 A3 A4 B

D E

I1 I2 I3

H K1

K2 J1 J2

Figure 7.—Morphology of Calobota angustifolia. A1–A4, leaves in abaxial view; A1, typical form; A2, Oran- jemund form; A3, short-leaved form; A4, broad-leaved form; B, flower in lateral view; C, standard petal; D, wing petal; E, keel petal; F, pistil; G, outer surface of the calyx (upper lobes to the left); H, androecium; I1, long, basifixed anther; I2, intermediate carinal anther; I3, short, dorsifixed anther; J1, bract; J2, bracteoles; K1, pod in lateral view; K2, pod in dorsal view. A1: Boatwright et al. 138, JRAU; A2: Ward 12435, WIND; A3: Giess & Robinson 13177, WIND; A4: Stirton & Zantovska 11426, NBG; B: Boat wright et al. 139, JRAU; C: Van Wyk 2847, JRAU; D–F: Boatwright et al. 139, JRAU; G: Lewis 4705, NBG; H, I: Boatwright et al. 139, JRAU; J: Whitehead 80.9.8, WIND; K: Boatwright et al. 138, JRAU. Scale bars: A & K, 10 mm; B–J, 1 mm. S T R E L I T Z I A 39 (2018) 27 brown, surface smooth (Figure 7). Flowering time: flowers for most of the year, January to November.

Distribution and habitat Calobota angustifolia is widely distributed along the West Coast of South Africa and Namibia (Figure 8). It forms an important component of Western Strandveld (FS 1 Lam- bert’s Bay Strandveld, FS 5 Langebaan Dune Strandveld; Rebelo et al. 2006), Richtersveld (SKr 3 Gariep Mountain Succulent Shrub- land and disturbed areas of SKr 13 Southern Richtersveld Scorpionstailveld; Mucina et al. Figure 8.—Distribution of Calobota angustifolia. 2006) and Namaqualand Sandveld (SKs 2 Northern Richtersveld Yellow Duneveld, SKs 4 Richtersveld Sandy Coastal Scorpionstail- pilose keel petals and the seeds are pale pink veld, SKs 5 Richtersveld Red Duneveld, SKs 6 with a smooth surface (C. lotononoides is a Oograbies Plains Sandy Grassland and SKs 13 small [up to 0.5 m], decumbent shrublet with Klawer Sandy Shrubland, the latter recorded unarmed inflorescences, extremely long wing as C. halenbergensis instead of C. angustifolia; petals, glabrous keel petals and the seeds are Mucina et al. 2006). Rebelo et al. (2006) re- brown with beige spots and a rugose surface). corded C. sericea as an important component It is also similar to C. halenbergensis, but dif- of FS 4 Saldanha Limestone Strandveld, but it fers in the sericeous leaves, pilose keel petals, is C. angustifolia that occurs in that area. hairy ovary and sericeous pods with up to 12 seeds (in C. halenbergensis the leaves are IUCN Red List category pilose, the keel petals glabrous and the ovary and pods glabrous with up to 7 seeds in the Considered of Least Concern (LC) by Rai- latter). mondo et al. (2009). On the Ecklon specimen (Ecklon s.n. in S), Discussion the type of C. angustifolia is represented by the two branches mounted at the bottom Calobota angustifolia is similar to C. lotonon right-hand side of the sheet – one with sever- oides, but differs in that it is an erect, divari- al flowers, the other with a single bud. Meyer cately branched shrub usually more than 1 m (1832) incorrectly cites the type locality as in height with inflorescences that sometimes ‘Uitenhage?’, but expresses doubt by placing terminate in a spine; the wing petals are al- a question mark after the locality. The speci- most as long as or slightly longer than the men in S comprises mixed elements, but the 28 S T R E L I T Z I A 39 (2018) two branches referred to above are accompa- long, bracteole 0.5–1.5 mm long. Flowers nied by a label in Meyer’s handwriting stating 9–11 mm long. Calyx 4.5–6.0 mm long; tube ‘Lebeckia angustifolia mihi’. The specimen 3.5–4.5 mm long; lobes 1–3 mm long. Stand Masson s.n. sub BM000794145 also compris- ard 9–13 mm long; claw 2.5–4.5 mm long; es mixed elements. The two specimens on the lamina 5.5–9.0 × (–3.5)4.5–10.0 mm. Wings left were collected by Masson and are marked 8.5–13.0(–14.0) mm long; claw 3.0–4.5(– with ‘1’ on the tape on the stems. The speci- 5.5) mm long; lamina 5.5–8.5 × 2.0–3.5 mm, men on the right was grown in the gardens at glabrous, with 5–8 rows of sculpturing. Keel Kew and has the note ‘Hort. Kew 1781’ asso- 8.0–12.5(–13.0) mm long; claw 3.5–4.5(– ciated with it. According to Stafleu and Cow- 5.5) mm long; lamina 4.5–7.5 × 2.5–4.5 mm, an (1976), most of the Aiton types are in the glabrous. Pistil ovary 5.5–9.5 × 0.5–1.1 mm Banks collection in BM. Drège 6474 ‘III, C’ with 11 to 20 ovules; style 3–5 mm long. Pods in P bears the type locality, shows the features sericeous, 18–35 × 2–3 mm. of the plant described by Meyer, has the characteristic Drège field labels and is therefore Distribution and habitat chosen as lectotype of Lebeckia multiflora. This form occurs around the Berg River, Drège s.n. ‘III, E, a’ in P displays the diagnos- Piquetberg, Clanwilliam, Velddrif, Lambert’s tic characters described by Meyer, bears the Bay, Vredenburg, Hopefield, Langebaan, type locality and is here chosen as lectotype Saldanha, Vanrhynsdorp, Heerenlogement, of Lebeckia decutiens var. a canescens. The Wallekraal, Hondeklipbaai, Garies and Klein- collection by Schinz (Schinz 811) is chosen see on the West Coast of South Africa and as lectotype of L. multiflora var. parviflora as does not extend into Namibia. It grows in it displays the characters of the variety and is well-drained sand, sandy loam, red sand or listed in the protologue. sand dunes between sea level and 500 m above.

Regional variation Diagnostic characters Four regional forms can be recognised based This form differs from the other forms in the on leaf morphology and geographical distri- long, narrow leaflets and relatively short pods. bution: The inflorescences are often spine-tipped. Populations of this form around Klawer and Form A (typical form) Vanrhynsdorp have very long wing petals and small flowers. Erect shrubs up to 1.8 m in height, often spinescent. Leaflets linear to narrowly oblanceo- Form B (broad-leaved form) late, sericeous, terminal 10–30 × 0.5–2.0(– 2.5) mm, lateral 8–23 × 0.5–2.0(–2.5) mm; Erect or spreading shrubs up to 2 m in petiole 11–35 mm long. Inflorescence height, sometimes spinescent. Leaflets obo- 27–90 mm long; with 10 to 27 flowers; vate to widely oblanceolate, sericeous, termi- pedicel 1–2 mm long; bract 1.5–2.0 mm nal 5–25 × 2–5 mm, lateral 5–25 × 2–4 mm; S T R E L I T Z I A 39 (2018) 29 petiole 8–30 mm long. Inflorescence 20– long; with 8 to 40 flowers; pedicel 1 mm long; 65 mm; with 7 to 18 flowers; pedicel 1–2 mm; bract 2–3 mm long, bracteole 1.5–2.0 mm bract 2 mm long, bracteole 1.0–1.5 mm long. long. Flowers 7–10 mm long. Calyx 4–6 mm Flowers 9–11 mm long. Calyx 5.0–6.5 mm long; tube 2.5–4.0 mm long; lobes 1.5– long; tube 3.5–4.0 mm long; lobes 1.0– 3.5 mm long. Standard 7.5–9.0 mm long; 2.5 mm long. Standard 10–11 mm long; claw claw 2–3 mm long; lamina 5.5–6.5 × 5.5– 3.0–3.5 mm long; lamina 6.5–8.0 × 7.5– 7.0 mm. Wings 8.5–10.0 mm long; claw 10.0 mm. Wings 10.5–11.5 mm long; claw 2.5–4.0 mm long; lamina 6–7 × 3–4 mm, 3–4 mm long; lamina 6.5–8.0 × 2.5–4.5 mm, glabrous, with 4–10 rows of sculpturing. glabrous, with 4–7 rows of sculpturing. Keel Keel 7.5–8.5 mm long; claw 2.5–3.5 mm 10–11 mm long; claw 3.5–4.0 mm long; lam- long; lamina semi-circular, 4–5 × 3.0– ina 6.5–8.0 × 3.5–5.0 mm, glabrous. Pistil 3.5 mm, glabrous. Pistil with ovary 5.5– with ovary 7.0–8.5 × 0.7–1.1 mm with 11 to 6.5 × 0.7–1.0 mm with 13 to 18 ovules; style 15 ovules; style 4.5–5.0 mm long. Pods seri- 3.0–3.5 mm long. Pods densely sericeous, ceous, 25–35 × 3 mm. 20–40 × 3–4 mm.

Distribution and habitat Distribution and habitat Occurs from the Saldanha area to Elands Bay, This form of Calobota angustifolia is restrict- Clanwilliam, Lambert’s Bay, Doring Bay, the ed to Namibia and occurs in the Lüderitz Bay mouth of the Olifants River, Brand-se-baai, area, including Boegoeberg, Bogenfels, Po- Garies, Wallekraal, Port Nolloth and Walvis mona, Chamais and Alexander Bay. It grows Bay along the West Coast of South Africa in well-drained or coarse sand, sometimes on and Namibia. It grows in deep limestone, cal- rocky soil or dolomite outcrops at altitudes crete sand or dunes at altitudes of between 4 of between 20 and 120 m. It is heavily grazed and 50 m and is grazed by livestock. by livestock.

Diagnostic characters Diagnostic characters This form differs from the others in the In this form the leaflets are very short with broad leaflets (always more than 2 mm wide) relatively long petioles. and relatively short inflorescences. Form D (Oranjemund form) Form C (short-leaved form) Erect shrubs up to 1.5(–1.8 m) in height. Erect or virgate shrubs up to 2 m in height. Leaflets narrowly oblanceolate, sericeous, Leaflets linear to oblanceolate, very densely terminal 5–14 × 0.5–1.0 mm, lateral sericeous, terminal 2–13 × 1–2 mm, lateral 5–9 × 0.5–1.0 mm; petiole 12–35 mm. In 2–7 × 0.5–1.0 mm; petiole relatively long, florescence 30–100 mm long; with 13 to 15 9–45 mm long. Inflorescence 30–200 mm flowers; pedicel 1–2 mm long; bract 1–3 mm 30 S T R E L I T Z I A 39 (2018) long, bracteole 1.0–1.5 mm long. Flowers (WIND); Lüderitz Bay (–CA), De Winter & Giess 7–10 mm long. Calyx 4.5–5.0 mm long; 6233 (BOL, K, PRE, WIND); Dinter 3838 (BM, BOL, K, NBG, PRE, S); Dinter 5996 (BM, BOL, K, NBG 2 tube 3.0–3.5 mm long; lobes 1–2 mm long. sheets, PRE); Giess & Van Vuuren 678 (PRE, WIND); Standard 6.5–8.0 mm long; claw 2–3 mm Lavranos & Pehleman 19641 (WIND 2 sheets); Leuen long; lamina 4.0–5.5 × 5.5–6.5 mm. Wings berger et al. 3306 (PRE, WIND); Merxmüller & Giess 7.0–8.5 mm long; claw 2.5–3.5 mm long; 28275 (K, PRE, WIND); Metz s.n. (WIND 2 sheets); lamina 4.5–5.5 × 3.0–3.5 mm, glabrous, Peter 47177 (K); Range 363 (BOL); Meyer 56 (WIND); with 8–12 rows of sculpturing. Keel 6.5– Volk 12825 (WIND); Wall s.n. (S); 13 mls [20.92 km] east of Lüderitz, road to Aus, Grasplatz (–CA), Giess 8.0 mm long; claw 2.5–3.5 mm long; lam- & Van Vuuren 738 (PRE, WIND); Lüderitz (–CA), ina 4.0–4.5 × 3.0–3.5 mm, glabrous. Pistil Hobart-Hampden 11 (BM); Lavranos & Barod 15409 with ovary 5–6 × 0.8–1.0 mm with 12 to (WIND); Moss 11462 (BM); Rogers 29588 (K 2 sheets); 19 ovules; style 2.5–3.5 mm long. Pods seri- just outside Lüderitz along main road (–CA), Man ceous, 20–32 × 2–3 mm. nheimer CM2015 (WIND); Angra Pequena (–CA), Marloth 4663 (NBG, PRE); Schenck 9 (PRE); Lüder- itz lagoon (–CA), Merxmüller & Giess 3133 (WIND); Distribution and habitat Lüderitz, just outside township (–CA), Müller & Jankowitz 275 (WIND); Essy Bay (–CA), Strohbach This form is centered in the Oranjemund 151 (WIND); Lüderitz Bay, Halbinsel (–CA), Walter district. It occurs in sandy areas, alluvial sand 56 (WIND); Wendt s.n. (WIND 2 sheets); Aus (–CB), and dunes near the mouth of the Orange Regius 23c (WIND); Grasplatz (–CB), Merxmüller & River and in the Oranjemund mining area at Giess 3052 (PRE, WIND); Venter 9160 (PRE). 2715 altitudes of between 10 and 100 m. (Bogenfels): Pomona (–AB), Dinter 6336 (BM, BOL, K, NBG 2 sheets, PRE); Kolle 13560b (NBG, PRE); Diagnostic characters Wall s.n. (S); Whitehead 80.9.8 (WIND); along road to Pomona (–AB), Mannheimer et al. CM898 (WIND); Leaflets of the Oranjemund form are short blue rocky ridge on way to Pomona town (–AB), and very narrow and the pods are also rela- Mannheimer CM 924 (WIND); Bogenfels (–AD), De tively short. Winter & Giess 6220 (K, PRE, WIND); Merxmüller & Giess 28326 (WIND); area and surrounds northeast of Bogenfels Arch (–AD), Powrie LWP1170 Additional specimens (WIND); Diamond Area 1 (–AD), Watmough 867 examined (PRE); Chamais (–DC), Merxmüller & Giess 28319 (K, PRE, WIND); Chamais valley in rock (–DC), NAMIBIA. 2214 (Swakopmund): Walvis Bay (–DC), Williamson 2647 (BOL); 3 km east of Chamais Bay (– Heywood s.n. (NBG). 2514 (Spencer Bay): ca. 2 km east DC), Williamson 4980 (WIND); Boegoeberg (–DD), of coast near East Hill between Oyster Cliffs and Easter Merxmüller & Giess 28304 (PRE, WIND); William point (–BD), Seely & Ward 54 (WIND); Spencer Bay son 2575 (BOL). 2816 (Oranjemund): between Oran- (–DB), Giess & Robinson 13177 (K, S, PRE, WIND); jemund and check point (–BC), Gess & Gess 97/98/68 Sydow 10179 (WIND). 2516 (Helmeringhausen): Hal- (J, WIND); Oranjemund (–CB), Merxmüller & Giess binsel (–CA), Walter 56 (WIND). 2615 (Lüderitz): 2267 (BM, K, WIND); sandy area near mouth of Or- Schurmhaudeberg (–AA), Hardy & Venter 4437 (K, ange River at Oranjemund (–CB), Metelerkamp 290 PRE, WIND); 14 mls [22.53 km] east of Lüderitz Bay (BOL); Schenck 238 (PRE); north of Orange River (– (–CA), Acocks 15654 (K, PRE 2 sheets); Lüderitz pen- CB), O’Callaghan et al. 5 (NBG, PRE); hills northeast insula (–CA), Clark & Müller 337 (WIND); De Win of Alexander Bay (–CB), O’Callaghan et al. 67 (NBG, ter & Hardy 7894 (K, WIND); Mannheimer CM2020 PRE); south bank of lagoon at mouth of Orange River S T R E L I T Z I A 39 (2018) 31

(–CB), Pillans 5636 (BOL); Oranjemund, near airport (Kamiesberg): Ezelkop, Garies (–AC), Scheffler 266 turnoff from main entrance road (–CB), Ward 11136 (NBG). 3117 (Lepelfontein): Brand-se-baai (–BD), De (K, PRE, WIND); Orange River, upper estuary (–CB), Villiers 34 (PRE, WIND); De Villiers 48, 49 (PRE); Ward 12435 (PRE); mining area 1 (–CB), Williamson Van Rooyen 2200 (PRE). 3118 (Vanrhynsdorp): 4844, 4851 (WIND); k122 mining area Ev5 control mouth of Olifants River, Papendorp (–CA), Arnold plot (–CB), Williamson 4884 (WIND). 916 (PRE); Vredendal, Olifants River Mouth (–CA), Le Roux & Ramsey 62 (NBG); Olifants River Mouth SOUTH AFRICA. 2816 (Oranjemund): Witbank (– (–CA), O’Callaghan 657 (NBG, PRE); Vredendal road DC), (BOL, K); 40 km from Port Nolloth Pillans 5207 (–CB), Steyn 464 (NBG); Doringbaai (–CC), Boucher on road to Alexander Bay (–DD), Germishuizen 4770 4054 (NBG, PRE); Hugo 2924 (PRE); Holrivier (– (PRE, WIND); Germishuizen 4771 (PRE); 32 km CB), Drège s.n. (S); between Vredendal and Lambert’s from Port Nolloth on road to Alexander Bay (–DD), Bay (–CD), Lewis 4705 (NBG); Schutte 269 (JRAU); Germishuizen 4813 (PRE). 2817 (Vioolsdrif): ‘ad ostia 10.7 km from Doringbaai to Donkinbaai (–CD), Stir fluvii Garip’ (–AA), Drège s.n. ‘III, B’ (P, K); sides of ton 6077 (K, PRE); Zandkraal (–DA), Acocks 14840 dry river bed at Kuboos (–AC), Pillans 5429 (BOL). (PRE); 11.7 km south of Redelinghuys (–DA), Acocks 2820 (Kakamas): Farm Harperspot, east of Riemvas- 24284 (K, PRE); Kys, southeast of Vredendal (–DA), maak (–AD), Gubb KMG13032 (WIND). 2916 (Port Bayer 6396 (PRE); Farm Liebendal, Vredendal (–DA), Nolloth): Port Nolloth (–BD), Bolus 9503 (K); Galpin Hall 3701 (NBG 2 sheets); Klawer (–DC), Andreae 474 & Pearson 7605 (BOL, K, NBG, PRE); Marloth 4747 (NBG, PRE); Heerenlogement (–DC), Esterhuysen (NBG, PRE); 1 km south of McDougalls Bay (–BD), 5547 (BOL); Vanrhynsdorp, foot of Nardousberge be- Raitt 302 (NBG); between Port Nolloth and Holgat low Witbakenkop on old Clanwilliam, Vanrhynsdorp (–BD), Schlechter s.n. sub STE 10850 (NBG 2 sheets); road (–DC), Hilton-Taylor 1613 (NBG); western as- Port Nolloth, at turn-off to McDougall Bay (–BD), pect of koppie on Vanrhynsdorp road, Klawer (–DC), Van Wyk 2847 (JRAU 2 sheets); 9 km east of Port Nol- Lavis 20236 (BOL, K 2 sheets); 6 mls [9.65 km] north loth (–BD), Zietsman & Zietsman 823 (PRE). 2917 of Klawer (–DC), Maguire 142 (NBG 2 sheets); 6 mls. (Springbok): sand by roadside between Anenous and [9.65 km] south of Klawer (–DC), Salter 5612 (BM, Chubiessis outspan (–BA), Pearson 5978 (BOL, K); BOL, K, PRE). 3217 (Vredenberg): Stompneus Point Oubeep Bay (–BD), Le Roux & Ramsey 144 (NBG); (–DB), Taylor 1515 (NBG); Brittania Bay (–DB), Tay 2 km south of Kleinsee (–CA), Hugo 2849 (PRE 2 lor 5189 (NBG, K, PRE); Farm Trekossenkraal (–DD), sheets); Buffels River mouth (–CA), Le Roux & Par Boucher 7065 (NBG); Cape Columbine Lighthouse sons 23 (PRE); Kleinsee (–CA), Theron 3858 (PRE); (–DD), Horrocks 182 (NBG 2 sheets); slopes south of between Brakwater and Komaggas (–DC), Compton Vredenburg (–DD), Hutchinson 247 (BM, BOL, K, 22071 (BOL, NBG). 3017 (Hondeklipbaai): 6 mls PRE); granite hills near Vredenburg (–DD), Marloth [9.65 km] southeast of Hondeklipbaai (–AD), Hall 7945 (PRE). 3218 (Clanwilliam): Farm Brakke Kuil 899 (BOL, NBG); Farm Oubees 339, 1.2 km east of 9, on road from Laaiplek to Elands Bay (–AB), Bösen Oubees/Wildepaardehoek boundary fence and 4.7 km berg & Rutherford 394 (NBG); Lamberts Bay (–AB), east of Springbok, Soebatsfontein road (–BA), Le Acocks 14182 (K, PRE), Clarke 633 (K, PRE); Henrici Roux & Lloyd 670 (NBG); between Brakwater and 3309 (PRE); Muir s.n. (NBG); Pole-Evans 26 (PRE); Komaggas (–BB), Compton 22071 (BOL, NBG); 3 mls Walsh s.n. (PRE); Groendam (–AB), Stirton 9349 [4.83 km] northwest of Wallekraal Post Office (–BC), (PRE); 5 km from Lamberts Bay to Elands Bay (–AB), Acocks 14933 (K, PRE 2 sheets); 5 mls [8.05 km] north- Stirton & Zantovska 11426 (JRAU, NBG, PRE); west of Wallekraal (–BC), Hall 895 (NBG 2 sheets); Wadrifsoutpan (–AB), Stirton 9340 (PRE); Farm Wallekraal, near Hondeklipbaai (–BC), Stirton 6036b Middelpos (–AB), Stirton 9387 (PRE); Nortier Ex- (K, PRE); 15 mls [24.14 km] east of Wallekraal on road perimental Station, Lambert’s Bay (–AB), Van Breda to Garies (–BC), Thompson 1090 (K, NBG); Groen 614 (NBG); Van Breda 4435 (PRE); 2 mls [3.22 km] River Mouth (–DC), Le Roux & Ramsey 276 (K, from Redelinghuys on Aurora road (–AD), Barker NBG, PRE); Le Roux & Ramsey 280 (NBG). 3018 9722 (NBG); Elands Bay (–AD), Barker 2627 (NBG), 32 S T R E L I T Z I A 39 (2018)

Britton 22 (NBG, PRE); Pillans 7962 (BOL); Taylor Mouth (–DA), Compton 11036 (NBG); Doorn River 3912 (NBG); Taylor 3915 (PRE); Walsh s.n. (PRE); (–DA), Schlechter 8059 (BM, BOL, K, PRE 2 sheets). Vredendal road from Graafwater (–BA), Barker 8545 3317 (Saldanha): 20 m off road in shooting range camp (NBG 3 sheets); between Leipoldtville and Graafwater parallel to dunes (–BB), Blake 91 (NBG); Hoedjies (–BA), Esterhuysen 3765 (NBG); Leipoldt 3765 (BOL, Bay (–BB), Bolus 12652 (PRE); Saldanha Bay (–BB), NBG); Nieuwoudtville turn-off after Graafwater on Grey s.n. (K); Marloth 10188 (PRE); koppie north of road to Lamberts Bay (–BA), Stirton 9197 (PRE); Saldanha Bay (–BB), Van Wyk 2696 (JRAU 5 sheets). Farm Rietfontein, Graafwater (–BA), Van Blerk 38 3318 (Cape Town): Lagoonside, Langebaan (–AA), (PRE); 5 mls [8.05 km] from Clanwilliam on Lam- Axelson 445 (NBG); roadside south of Langebaan vil- berts Bay road (–BB), Gillett 4045 (PRE); Rocher Pan lage (–AA), Boucher 2811a (NBG); near Langebaan Nature Reserve (–CB), Heyl 44 (NBG, PRE); Le Roux (–AA), Chaplin s.n. (NBG); Marloth 13176 (PRE); & Van Rooyen 1 (NBG); on road to Hopefield from Geelbek (–AA), Compton 19906 (NBG); Langebaan Velddrif (–CC), Boatwright et al. 138 (JRAU); road Peninsula above Preekstoel (–AA), O’Callaghan & from Velddrif to Saldanha Bay (–CC), Boatwright et Steensma 1650 (NBG); Hopefield (–AB), Bachmann al. 139 (JRAU); Velddrif (–CC), Compton 15940 1889 (BM, K); 10.7 mls [17.22 km] from Hopefield (NBG); Goldblatt 6016 (PRE); Lewis 899 (NBG); to Vredenberg (–AB), Marsh 182 (NBG, PRE); 14 km Mauve 5356 (K, PRE); 1 km from road between Veld- from Hopefield to Langebaanweg (–AB), Stirton drif and Vredenburg on road to Stompneusbaai (– 10709 (K); Schrywers Hoek (–AD), Macnac 1020 CC), Grobbelaar 2546 (PRE); Velddrif, next to road (PRE); near Bergrivier and Zwartland (–BD), Ecklon before the bridge going over the Berg River (–CC), Le & Zeyher 1346 (S); sandy hill slopes behind the vil- Roux 3 (JRAU 4 sheets); between Hopefield and Pa- lage Langebaan (–DC), Leighton s.n. sub BOL 32422 ternoster (–CC), Leipoldt 3764 (BOL); 1 ml [1.61 km] (BOL). 3322 (Oudtshoorn): Waenskloof, Cango from Velddrif, on banks of Berg River (–CC), Marsch Valley (–AC), Moffett 494 (PRE). Precise local- 1265 (K, NBG, PRE); ca. 6 km east of Velddrif, near ity unknown: Olifantsrivier, Drège s.n. (P); without Ouwerf (–CC), O’Callaghan 1237 (NBG); Berg River locality, Galpin & Pearson 7579 (PRE); Leipoldt s.n. Station (–CD), Barker 4050 (NBG); railway enclosure (NBG); Range 1941 (NBG); Bokkeveld Mountains, along road 0.5 mls [0.8 km] from Berg River Station Leipoldt 730 (PRE); Little Namaqualand, Krapohl (–CD), Boucher 83 (NBG, PRE); South side of Berg 11142 (PRE); Lus district, Merxmüller & Giess 2349 River (–CD), Leighton 614 (BOL); Piquetberg (–DA), (WIND); sandy ravine below Doornpoort, Pearson Van Aarde s.n. (NBG); Kapteinskloof (–DC), Stirton 6017 (BOL, K); Waterkloof at Doornpoort, Pillans 6132b (K, PRE). 3219 (Wuppertal): Doorn River 5377 (BOL); Namaqualand, Scully s.n. (NBG). S T R E L I T Z I A 39 (2018) 33

3. Calobota cinerea (E.Mey.) Boatwr. & B.-E.van Wyk, S. African J. Bot. 75: 553. 2009. Bas- ionym: Lebeckia cinerea E.Mey., Comm. Pl. Afr. Austral. [Meyer] 1: 35. Feb. 1836. TYPE: SOUTH AFRICA, hills near Noagas [grid unknown], Drège s.n ‘III, B’ (lectotype, designated here, P!, flowering specimen on the left; isolectotypes, BM!, K!, P!). Lebeckia cinerea var. schäferi Dinter, Plantae novae Schäferianae: 342. 1920. TYPE: NA- MIBIA, ‘Granitberg zwischen Bogenfels und Prinzenbucht’ [2715 AD], Schäfer 598 (B†?, specimen not located).

Erect, multi-stemmed, unarmed shrub up ovary linear, 6–8 × 0.8–1.2 mm with 10 to 16 to 1.5 m in height. Branches green; young ovules; style shorter than ovary, 5.0–8.5 mm branches sericeous or silky to tomentose; long. Pods linear, terete, subsessile to shortly older branches sericeous or silky with light stipitate, 15–40 × 2–4 mm, ±5- to 6-seeded, brown bark. Leaves digitately trifoliolate, tomentose, dehiscent; endocarp hairy. Seeds deciduous; petiole 3–14 mm long, longer oblong-reniform to reniform, 2–3 × 1.5– or shorter than leaflets; leaflets widely obo- 2.0 mm, mature seeds light brown to orange, vate to elliptic or oblanceolate, alternate, often mottled with brown, surface smooth sericeous on both surfaces, subsessile, ter- (Figure 9). Flowering time: mainly flowering minal leaflet 5–21 × 1.5–6.0 mm, lateral in spring (August to November) and fruiting leaflets 3–17 × 1.5–5.0 mm; apex recurved- within the same period. Some specimens col- mucronulate to acute; base cuneate. Inflores lected in Namibia were flowering and fruit- cence 30–200 mm long, racemose, with 5 to 30 ing in April and May. flowers; pedicel 2–3 mm long; bract 1–4 mm long, linear, sericeous; bracteoles 1–4 mm Distribution and habitat long, linear, sericeous. Flowers 11–16 mm long, pale to bright yellow. Calyx 5–8 mm Calobota cinerea occurs from southwestern long, sericeous; tube 3.5–5.0 mm long; Namibia southwards to South Africa as far lobes 1.5–3.5 mm long, deltoid. Standard south as Clanwilliam (Figure 10). It occurs 11.0–13.5 mm long; claw linear, 3.5–4.5 mm on well-drained, sandy, loamy soil or red long; lamina ovate, 8–10 × 7–11 mm; apex sand often next to rivers, at altitudes of be- obtuse; dorsal surface densely pubescent. tween ca. 100 and 500 m. It is recorded to Wings 9–13 mm long; claw 2.5–3.5 mm be grazed by livestock or gemsbok. Accord- long; lamina oblong, shorter than keel, 5.5– ing to Mucina et al. (2006), C. cinerea is an 8.5 × 3–4 mm, glabrous or rarely pilose, with important component of Richtersveld (SKr 5–8 rows of sculpturing. Keel 11–14 mm 6 Stinkfontein Eastern Apron Shrubland) long; claw 3.5–4.5 mm long; lamina boat- and Namaqualand Sandveld (SKs 2 North- shaped, 7.5–10.0 × 4.5–5.5 mm, pilose. Pis ern Richtersveld Yellow Duneveld, SKs 5 til subsessile to shortly stipitate, pubescent; Richtersveld Red Duneveld, Sks 6 Oograbies 34 S T R E L I T Z I A 39 (2018)

A1 A2 B1 B2

D F

I1 I2 I3 G H

K1 J

Figure 9.—Morphology of Calobota cinerea. A1, A2, leaves in abaxial view; B1, bract; B2, bracteoles; C, flower in lateral view; D, standard petal; E, outer surface of the calyx (upper lobes to the left); F, wing petal; G, keel petal; H, androecium; I1, long, basifixed anther; I2, intermediate carinal anther; I3, short, dorsifixed anther; J, pistil; K1, pod in lateral view; K2, pod in dorsal view. A1: Boatwright et al. 136, JRAU; A2–K: Boatwright et al. 150, JRAU. Scale bars: A & K, 10 mm; B–J, 1 mm. S T R E L I T Z I A 39 (2018) 35

Plains Sandy Grassland, SKs 8 Namaqualand Coastal Duneveld).

IUCN Red List category Considered of Least Concern (LC) by Rai- mondo et al. (2009).

Discussion Calobota cinerea is similar to C. linearifolia, but differs in the low, spreading habit as opposed to the virgate or erect habit of C. line arifolia. The leaves are trifoliolate and obovate to elliptic or oblanceolate as opposed to Figure 10.—Distribution of Calobota cinerea. the simple linear to oblanceolate or spathulate leaves of C. linearifolia. In C. cinerea the inflorescences are more densely flowered Pomona (–AB), Dinter 6346 (BM 2 sheets, BOL, K, with up to 30 flowers (up to nine flowers in NBG 2 sheets, PRE); Wall s.n. (S); Pomona area, west C. linearifolia), the pods are tomentose and facing slope ±15 km from houses going east (–AB), Mannheimer et al. CM889 (WIND); Prince of Wales not constricted between the seeds, and the Bay (–AB), Marloth 5248 (PRE); Granite Mountain seeds light brown to orange or mottled with (Granietberg) near Bogenfels (–AD), Schäfer 13 (NBG). brown (pods sericeous, constricted between 2716 (Witputz): Diamond Area 1, Inselberg, at beacon the seeds and the seeds light brown or grey in 992, end of Uguchab River (–CA), Germishuizen 10170 C. linearifolia). (PRE). 2815 (Diamond Area): Klinghardt Mountains (–BB), Dinter 3918 (BM, BOL, NBG, PRE, S). The Drège specimen in P was annotated by SOUTH AFRICA. 2816 (Oranjemund): ±40 km Meyer himself and is here chosen as lectotype. from Brandkaros (red dune area) near farm fence (– BC), Petersen s.n. (NBG); Oranjemund (–CB), Clark 8 (WIND); sandy depressions north of Witbank (–DA), Additional specimens Pillans 5221 (BOL, K); 38 mls [61.14 km] north of examined Port Nolloth on way to Alexander Bay (–DA), Werger 517 (K, PRE); Goariepvlakte (–DB), Jürgens 22334 NAMIBIA. 2518 (Tses): Kovisberge (–BB), Wall 2 (PRE); 35 km from Port Nolloth on road to Alexander (S). 2615 (Lüderitz): Angra Pequena (–BA), Marloth Bay at Holgat River (–DD), Germishuizen 4772 (PRE, 4809 (PRE); Marloth 14109 (K); Kovisberge (–CB), WIND); Holgat (–DD), Jürgens 22734, 28804 (PRE). Dinter 6292 (BM, BOL, K, NBG); Halenberg (–CB), 2820 (Kakamas): Between Augrabies and Anenous (– Merxmüller & Giess 3089 (WIND). 2616 (Aus): near CB), Bolus 428 (BM, BOL, K, NBG, PRE); Bolus 6548 Aris on the Gariep (–BD), Drège s.n. ‘III, B’ (BM, P); (BOL, K). 2917 (Springbok): Anenous flats, ca. 8 km Farm Klein-Aus (–CA), Merxmüller & Giess 2998 west of Farm Grasvlakte (–AB), Goldblatt & Man (PRE, WIND); 15 mls [24.14 km] west of Aus (–CB), ning 9461 (K, NBG); on road to Port Nolloth from Giess & Van Vuuren 833 (K, WIND). 2715 (Bogenfels): Steinkopf (–AC), Boatwright et al. 150 (JRAU); sandy 36 S T R E L I T Z I A 39 (2018)

slope 40 km east of Port Nolloth (–AC), Goldblatt & Hall 3700 (NBG 2 sheets); between Vredendal and Manning 9289 (NBG); 15 mls [24.14 km] north of Lutzville (–DA), Lückhoff s.n. (NBG); 53 km from Port Nolloth (–AC), Herre s.n. (NBG); between Kaus, Clanwilliam to Vanrhynsdorp (–DA), Stirton 5944 Natvoet and Doornpoort (–AD), Drège s.n. (S); Farm (K, PRE); Farm Spes Bona, Inedendahl (–DC), De Nuwefontein 147 northwest of windmill at 28th mile partment of Agriculture 29 (PRE); Klawer (–DC), (–DB), Le Roux 4038 (NBG). 3017 (Hondeklipbaai): Henrici 3344 (PRE); western aspect of koppie, Van- Taaibosvlei in dunes (–BC), Marloth 12418 (NBG, rhynsdorp road near Klawer (–DC), Lavis 20235 PRE). 3118 (Vanrhynsdorp): Hardeveld southwest of (BOL). 3218 (Clanwilliam): 8 km from Clanwilliam Nuwerus (–AB), Marloth 8233 (NBG, PRE); sand- on road to Vanrhynsdorp (–BB), Boatwright et al. dunes near Doorn River (–BB), Marloth 2637 (NBG, 136, 210 (JRAU); 5 km north of Clanwilliam (–BB), PRE); Farm Quaggaskop, 16 km on Douse the Glim Grobbelaar 2004 (PRE); 10 km before Clanwilliam turn-off, 24 km north of Vanrhynsdorp (–BC), Le from Vanrhynsdorp (–BB), Van Wyk 2598 (JRAU 2 Roux 2066 (NBG); towards ‘Douse the Glim’, Kners sheets, PRE). Precise locality unknown: without lo- vlakte (–BD), Stirton 9373 (PRE); Lutzville (–CB), cality, Drège s.n. (K); Sandveld, Richtersveld, Herre s.n. Stirton 6062 (PRE); Vredendal road (–DA), Barker sub STE 11879 (NBG); Andaus Poort, Marloth 12250 5692 (BOL, NBG 2 sheets); Farm Liebendal (–DA), (NBG, PRE). S T R E L I T Z I A 39 (2018) 37

4. Calobota cuspidosa (Burch.) Boatwr. & B.-E.van Wyk, S. African J. Bot. 75: 553. 2009. Basionym: Spartium cuspidosum Burch., Trav. S. Africa 1: 348. 1822. Genista cuspidosa (Burch.) DC., Prodr. [DC.] 2: 147. 1825. Lebeckia cuspidosa (Burch.) Skeels, U. S. D. A. Bur. Pl. Industr. Invent. Seeds 233: 38. 1911; Lebeckia cuspidosa (Burch.) Druce, Rep. Bot. Exch. Club Brit. Isles 1916: 631. 1917., nom illeg. TYPE: SOUTH AFRICA, Betw. Gati- kamma & Klaarwater [now Griquatown; 2823 CC], Burchell 1697 (holotype, K!). Lebeckia macrantha Harv. in Harv. & Sond., Fl. Cap. [Harvey] 2: 83–84. 1862; Wilman, Preliminary checklist of the flowering plants and ferns of Griqualand West (southern Africa): 53. 1946. TYPE: SOUTH AFRICA, ‘Zooloo country’ [grid unknown], Miss Owen s.n. (holotype, TCD, photo!).

Erect, multi-stemmed spinescent shrub up glabrous, with 5 rows sculpturing. Keel to 4 m tall. Branches green; young branches 13.5–19.0 mm long; claw (3.5–)5.0–7.0 mm sparsely pubescent; older branches pubes- long; lamina oblong, 10.0–12.5 × 5–6 mm, cent to glabrous with light brown bark. glabrous. Pistil shortly stipitate, glabrous; Leaves unifoliolate or rarely trifoliolate, de- ovary linear, 7–14 × 1.0–1.6 mm with 9 to 11 ciduous; petiole short, 1–2 mm long (up ovules; style shorter than ovary, 5.5–9.0 mm to 5 mm long when trifoliolate); alternate, long. Pods oblong to slightly falcate, laterally upper and lower surfaces pilose, linear to compressed, shortly stipitate, (21–)28–49(– slightly obovate, 6–9(–15) × 2–3(–6) mm; 53) × (5–)6–10 mm, ±4- to 6-seeded, gla- apex obtuse to shallowly retuse; base attenu- brous, indehiscent; endocarp glabrous. Seeds ate. Inflorescences (25–)40–60(–90) mm reniform, 3–4 × 2.5–3.0 mm, mature seeds long, racemose, with (4–)6 to 12(–24) flow- light brown, surface smooth (Figure 11). ers; pedicel 2–4(–5) mm long; bract 0.9– Flowering time: this species flowers mainly 1.2 mm long, triangular to slightly linear; in spring and summer (October to January), bracteoles 0.5–0.8(–1.0) mm long, slightly but flowering also sometimes occurs in win- ovate. Flowers (15–)16–18(–20) mm long, ter (April to May). bright yellow. Calyx (3.8–)4.4–5.4 mm long, pubescent on the outer surface; tube Distribution and habitat (2.5–)3.0–4.0 mm long; lobes narrowly triangular, (1.2–)1.8–2.4(–2.9) mm long, del- Calobota cuspidosa occurs in the North- toid. Standard 11.0–18.5 mm long; claw ern Cape Province and is centred around linear, 3–6 mm long; lamina ovate, 8.5– the towns of Kuruman and Griquatown 12.5 × (7.5–)8.0–10.0 mm, glabrous; apex at altitudes of between 975 and 1 700 m. obtuse. Wings 10–16 mm long; claw (3–)5– It is mainly found in rocky, loamy or sandy 6 mm long; lamina oblong, shorter than the soils (Figure 12). According to Rutherford keel, (7.5–)9.0–10.5 × (3.0–)4.0–5.5 mm, et al. (2006) this species is an important, 38 S T R E L I T Z I A 39 (2018)

A1 A2

B C

E F

G I1 I2 I3

J1 J2 K1

Figure 11.—Morphology of Calobota cuspidosa. A1, A2, leaves in abaxial view; B, outer surface of the calyx (upper lobes to the left); C, flower in lateral view; D, standard petal; E, wing petal; F, keel petal; G, pistil; H, androecium; I1, long, basifixed anther; I2, intermediate carinal anther; I3, short, dorsifixed anther; J1, bract; J2, bracteoles; K1, pod in lateral view; K2, pod in dorsal view. A1: MacDonald 76/3, NBG; A2: Ethel Anderson 591, BOL; B: Gubb 12584, PRE; C–H: Van Wyk 3055, JRAU; I: Gubb 12584, PRE; J: Van Wyk 3055, JRAU; K: Boatwright et al. 92, JRAU. Scale bars: A & K, 10 mm; B–J, 1 mm. S T R E L I T Z I A 39 (2018) 39 prominent component of Kuruman Moun- tain Bushveld (SVk 10 Eastern Kalahari Bushveld) where it is common in low, open bushveld.

IUCN Red List category Considered of Least Concern (LC) by Rai- mondo et al. (2009).

Discussion Calobota cuspidosa is similar to C. psiloloba, but the large flowers, pods and seeds are dis- Figure 12.—Distribution of C. cuspidosa. tinctive for the former (in C. psiloloba the flowers, pods and seeds are much smaller). Calobota cuspidosa reaches heights of up to 4 m, while C. psiloloba only reach heights of sub TRV 31766 (PRE); On road to Olifantshoek (– AD), Germishuizen 5614 (BOL, PRE); 3 mls [4.8 km] 1.2 m. The young stems in this species are east of Kuruman (–AD), Grobbelaar 1112 (PRE); Ku- normally densely tomentose and the older ruman Municipal Reserve (–AD), MacDonald 76/3 (J, ones remain at least slightly pubescent (in NBG, PRE); Buta Asbestos Mine (–AD), Peeters, Ger C. psiloloba the branches are glabrous or only icke & Burelli 264 (J, PRE); ca. 12 km from Kuruman to slightly pubescent). Vryburg (–AD), Van Wyk 3053 (JRAU); 11 km from Kuruman to Vryburg (–AD), Van Wyk 3055 (JRAU); Kuruman Hills (–AD), Verdoorn & Dyer 1763 (PRE, The specimen Miss Owen s.n. in TCD is the K); Vredebron, Blikfontein (–CA), Gubb KMG 10989 only one cited by Harvey in the original de- (PRE); 10 km west of Kuruman on road to Upington scription of Lebeckia macrantha and was also (–CA), Van Wyk BSA1241 (PRE, WIND); 30 km from seen by him as is indicated by the exclamation Kuruman to Billingshurst (–CD), Arnold & Musil 500 mark following the specimen citation in Har- (PRE); Hillsides from Griquatown to Khosis (–CD), Bryant J379 (PRE); 30 km southeast of Kuruman (– vey (1862). It must therefore be the holotype DA), Jordaan CBK 26 (PRE); Road from Reivilo to of L. macrantha despite the incorrect locality. Kuruman, farm Mattana (–DB), Joffe 650 (PRE). 2724 (Taung): Blesmanspost (–CA), Burtt-Davy 9665 (K). 2820 (Kakamas): Cape Plateau, Baviaans ‘Kloof’ [= Additional specimens Baviaans Krantz], (–DA), Hutchinson 3001 (BOL, K, examined PRE). 2822 (Glen Lyon): Hills at Vaalwater (–DA), Acocks 2455 (BOL 2 sheets, K 2 sheets); between Gri- SOUTH AFRICA. 2723 (Kuruman): Batlaros, Bosjes- quatown and Volop, Bakenkop, north of tarred road (– mans doorn (–AC), Silke 16785 (BOL); ca. 10 km DB), Gubb KMG 10988 (PRE). 2823 (Griquatown): west of Kuruman on the road to Olifantshoek (–AD), Postmasburg (–AC), Wilman s.n. (K); 19.5 km from Dlamini, Nkuna & Van Wyk EvW577 (K); Kuruman Griquatown to Campbell (–AD), Boatwright et al. (–AD); Pole-Evans 2108, 2480 (K, PRE); Van Son s.n. 92 (JRAU); between Postmasburg and Griquatown 40 S T R E L I T Z I A 39 (2018)

(–CA), Hutchinson 3039 (BM, BOL, K, PRE); Jasper Schmidtsdrift, Ruigtefontein (–CA), Gubb KMG veld (–CB), Wilman 4685 (K); Griquatown common- 12584 (PRE); 9 mls [14.5 km] west of Schmidtsdrift age, the Asbestos Hills (–CC), Acocks H1049 (PRE); (–CA), Leistner & Joynt 2690 (K, PRE); Schmidtsdrif Griquatown (–CC), Barret-Hamilton s.n. (BM); Botha (–CA), Wall s.n. (S); Barkly West, Tierfontein (–DA), 2996 (PRE); Burchell 1697, Pole-Evans 22 (K, PRE); 1 Acocks 719 (K 2 sheets); 14 mls [22.5 km] from Vryburg ml [1.6 km] east of Griquatown (–CC), Leistner 890 on Schweizer Reneke road (–DD), Carr 5 (PRE). 2922 (K, PRE); ca. 5 km from Griquatown on R64 on the (Prieska): 6 mls [9.6 km] west of Abrahmsdam (–BA), road to Upington (–CC), Van Wyk 2534 (JRAU 3 Codd 1263 (K, PRE). 2923 (Douglas): Mazelsfontein sheets); Broken Hill, north of Campbell (–DA), Ger Griqualand West (–BA), Anderson 591 (BOL, PRE); mishuizen 8523 (PRE). 2824 (Kimberley): Koopmans- near Douglas (–BB), Kotze 783 (PRE). Precise locality fontein, Farm Geluk (–AA), Hanekom 2214 (K, PRE); unknown: Namaland, Marloth 1042 (PRE, 2 sheets). S T R E L I T Z I A 39 (2018) 41

5. Calobota cytisoides (Bergius) Eckl. & Zeyh., Enum. Pl. Afric. Austral. [Ecklon & Zeyher] 2: 191 Jan. 1836. Basionym: Spartium cytisoides Bergius, Descr. Pl.: 199. 1767; L.f., Suppl. Pl. 320. 1781. Lebeckia cytisoides (Bergius) Thunb., Nov. Gen. Pl. [Thunberg]: 143. 1800. TYPE: SOUTH AFRICA, e Cap. B. Spei [grid unknown], Grubb s.n. sub Bergius 236.57 (holotype, SBT!). Ebenus capensis L., Mant II: 264. 1771. Cytisus capensis Lam., Encycl. [Lamarck] 2: 249 (1786); Lebeckia capensis (L.) Druce, Rep. Bot. Exch. Club Brit. Isles 1916: 631. 1917. TYPE: not seen. Crotalaria pulchella Andrews, Bot. Repos. (dup) 6: plate 417 (Jan–Mar 1805). Lebeckia pulchella (Andrews) Walp., Linnaea 13: 477 (1839). Calobota pulchella (Andrews) Eckl. & Zeyh., Enum. Pl. Afric. Austral. [Ecklon & Zeyher] 2: 191 (Jan. 1836). TYPE: Andr. Bot. Repos. (dup) 6: plate 417. 1805. (iconotype). ?Lebeckia marginata E.Mey., Comm. Pl. Afr. Austral. [Meyer] 1: 35. Feb.1836. TYPE: without locality, Drège s.n. (S!).

Erect, diffuse, multi-stemmed, unarmed basal callosities, 18.5–27.0 mm long; claw shrub up to 2 m in height. Branches green; linear, 4–9 mm long; lamina widely ovate, young branches sericeous; older branches seri- 13–19 × 14.5–26.5 mm; sericeous along ceous to pilose with brown bark. Leaves digi- dorsal midrib and apex; apex obtuse. Wings tately trifoliolate to rarely 5-foliolate, persis- 18–28 mm long; claw 5.5–9.0 mm long; lam- tent; petiole (8–)10–40 mm long, as long as or ina oblong to ovate, shorter or ± as long as shorter than leaflets; leaflets elliptic to oblan- keel, 12.5–17.5 × 7–12 mm, glabrous, with ceolate, alternate, sericeous, subsessile, ter- 9–16 rows of sculpturing; apex obtuse. Keel minal leaflet (12–)14–55 × 3–10(–17) mm, 19.0–29.5 mm long; claw 6.0–12.5 mm long; lateral leaflets (11–)13–45 × 3–9(–15) mm, lamina boat-shaped, 13–20 × 6–10 mm, seri- apex mucronulate to somewhat truncate, ceous on terminal parts; apex obtuse. Pistil base angustate. Inflorescence 70–255(– subsessile to shortly stipitate, glabrous; ovary 300) mm long, racemose, with 7 to 25 flowers; linear, 14–22 × 1.2–2.0 mm with 20 to 34 pedicel relatively long, 3–11 mm long; bract ovules; style shorter than ovary, 7.5–13.5 mm 3–6 mm long, elliptic to ovate, pubescent on long, glabrous. Pods linear, sometimes some- outer surface; bracteoles 1–4(–5) mm long, what clavate, terete, subsessile to shortly narrowly ovate to linear, pubescent on outer stipitate, 45–70 × 3–5 mm, ±5- to 18-seeded, surface. Flowers 18–35 mm long, bright yel- glabrous, dehiscent; endocarp hairy. Seeds low. Calyx 9–13 mm long, glabrous or very oblong-reniform, 2.8–3.5 × 2.0–2.6 mm, rarely pilose; tube 6.5–10.5 mm long; lobes mature seeds light brown, surface smooth 1.5–4.0 mm long, deltoid. Standard with (Figure 13). Flowering time: mainly flowering 42 S T R E L I T Z I A 39 (2018)

C A B

D E

F G H

I1 I2 I3

K1 K2 L1

K3 K4 K5 L2

Figure 13.—Morphology of Calobota cytisoides. A, B, leaves in abaxial view; C, flower in lateral view; D, outer surface of the calyx (upper lobes to the left); E, standard petal; F, wing petal; G, keel petal; H, androecium; I1, long, basifixed anther; I2, intermediate carinal anther; I3, short, dorsifixed anther; J, pistil; K1, K2, bracts; K3–K5, bracteoles; L1, pod in lateral view; L2, pod in dorsal view. A: Van Wyk 3117, JRAU; B: Le Roux 2, JRAU; C–K1, K4, K5: Boatwright et al. 114, JRAU; K2, K3: Barker 9694, NBG; L: Van Wyk 2439, JRAU. Scale bars: A, B & L, 10 mm; C–J, 1 mm. S T R E L I T Z I A 39 (2018) 43

component of Shale Fynbos (FFh 8 Mon- tagu Shale Fynbos), Shale Renosterveld (FRs 9 Swartland Shale Renosterveld, listed as C. cinerea), and Rainshadow Valley Karoo (SKv 7 Robertson Karoo).

IUCN Red List category Considered of Least Concern (LC) by Rai- mondo et al. (2009).

Discussion Calobota cytisoides is similar to C. sericea, but Figure 14.—Distribution of Calobota cytisoides. differs in the mucronulate leaf apices, long pedicels, larger flowers, sometimes ovate bracts and bracteoles, glabrous or pilose calyx, and fruiting in late winter to spring (June to larger and glabrous pistil and pods (in C. seri November), one flowering specimen was col- cea the leaf apices are acute, the pedicels short- lected in April. er, the flowers smaller, the bracts and bracteoles always linear, the calyx sericeous and the pistil and pods shorter and pubescent). Distribution and habitat According to Stafleu & Cowan (1976), Calobota cytisoides is a common and widely Henry Andrews was a botanical painter and distributed species. It can be found from engraver. His artworks were based on living Oudtshoorn through the Western Cape to specimens, but no herbarium specimens are around Montagu and Worcester and north- known to exist. Due to the fact that the paint- wards through Wupperthal and Clanwilliam ing in Botanist’s Repository for New and Rare as far as Paardekraal north of Nieuwoudtville Plants, London (dup) 6 (plate 417) leaves no (Figure 14). It has a more southern distribu- doubt as to the identity of the species, we here tion than the closely related C. sericea and choose it as iconotype of Crotalaria pulchella. has been collected at altitudes of between 20 Lebeckia marginata was described by Meyer and 1 000 m in renosterveld vegetation on (1836) without listing locality information. sandstone, shale, well-drained sand and clay. The only specimen located is in S and is ster- Calobota cytisoides is often found on western, ile, but the mucronulate apices of the leaflets southern or southwestern slopes, along riv- look similar to those of Calobota cytisoides. ers or in riverbeds and also along disturbed This taxon might merely represent a synonym roadsides. Rebelo et al. (2006) and Mucina of the latter as suggested by Ecklon and Zey- et al. (2006) list this species as an important her (1836), but its identity remains unclear. 44 S T R E L I T Z I A 39 (2018)

Additional specimens River Valley, 23 km north of Citrusdal (–BD), Hugo 426 (K, NBG, PRE); Rondegat River, 10 km from Algeria examined to Clanwilliam (–BD), Stirton 5926a (K, PRE); roadside between Citrusdal and Clanwilliam (–BD), Taylor SOUTH AFRICA. 3118 (Vanrhynsdorp): ‘Pillar se 10630 (NBG, PRE); Van Wyk 2313 (JRAU 2 sheets); punt’, northern point of Bokkeveld Mountains, plateau halfway between Clanwilliam and Citrusdal (–BD), on farm Paardekraal (–BB), Van Wyk 1248 (NBG, Van Wyk 2439 (JRAU 2 sheets); Van Wyk 2441 PRE, UPS); Vleikraal, east of Klawer (–DA), Walters (JRAU); Olifants River Valley between Klawer and Cit- 178 (K, NBG); 17.2 km from caravan park, Clanwilliam rusdal (–BD), Wilman 828 (BOL); St Helena Bay, on old road to Citrusdal (–DC), Gess 91/92/1 (GRA); Steenberg Cove (–CA), Wisaru 1710 (NBG); on road 23.9 km from Clanwilliam (–DC), Gess 91/92/10 between Misverstand Dam and Velddrif (–CC), Le (GRA); Klawer (–DC), Lavis 29233 (BOL); 29 km Roux 2 (JRAU 2 sheets); 9 mls [14.48 km] from Veld- from Clanwilliam to Vanrhynsdorp (–DC), Stirton drif to St. Helena Bay (–CC), Marsch 1272 (K, NBG); 5943 (K, PRE). 3119 (Calvinia): Vanrhyns Pass (–AC), 2 mls [3.22 km] north-west of Sauer Post Office (–CD), Acocks 14717 (K, PRE 2 sheets); Bond 1147 (NBG); Es Acocks 14534 (K, PRE 3 sheets); Moutonsvlei, west of terhuysen 5312 (BOL); Schutte 286 (JRAU 2 sheets); Piquetberg Mountain complex (–DA), Goldblatt 7197 Van Wyk 3117 (JRAU); Skuinshoogte Pass (–AC), Nie (NBG, PRE, S); base of north slopes below nek at head mand 1 (JRAU); Niewoudtville, Oorlogskloof Nature of Kapteinskloof, Piquetberg (–DA), Pillans 7701 Reserve (–AC), Pretorius 105 (NBG, PRE, WIND); (BOL); Het Kruis (–DA), Stephens & Glover 8686 (BM, Nieuwoudtville (–AC), Schmidt 636 (K); Meulsteenvlei NBG 2 sheets, K, PRE); Kapteinskloof (–DA), Stirton near Nieuwoudtville (–AC), Taylor 3942 (NBG); Lo- 6133a (K, PRE); near Warmbad (–DB), Hutchinson kenberg, Stinkfontein Hills (–CA), Leistner 451 (NBG, 1132 (BM, BOL, K, PRE); Olifants River Valley (–DB), PRE). 3217 (Vredenburg): Vredenburg (–DB), Grob Marloth 7121 (PRE); Piekenierskloof Pass (–DB), Van belaar 2549 (PRE); above Stompneus (–DB), Booysen Wyk 3219 (JRAU); 3 km south of Piquetberg (–DC), 141 (NBG); 1 ml [1.61 km] east of Paternoster (–DD), Ihlenfeldt 1023 (PRE); 3 mls [4.83 km] from Piquet- Barker 9694 (NBG 2 sheets); Kasteelberg (–DD), Strid berg up Versfeld Pass (–DC), Thompson 762 (K, NBG, & Strid 38078 (NBG). 3218 (Clanwilliam): Graafwa- PRE); outside Piquetberg (–DC), Thorne s.n. (NBG); ter (–BA), Compton 24209 (NBG); Bulshoek Barrage, Piquetberg (–DD), Bolus 13536 (BOL, PRE 2 sheets); Olifants River Valley (–BB), Barker 6426 (NBG 2 Grant 4714 (PRE); Guthrie 2584 (NBG); Schlechter sheets); Clanwilliam (–BB), Bayliss BRI593 (GRA, K, 5233 (BM, BOL, K); Schlechter s.n. sub TRV 3377 PRE, S); Davidson s.n. (J); Ecklon & Zeyher 17 (NBG); (PRE); roadside near Piquetberg (–DD), Hafström & Eliovson 161 (J); Henrici 2105 (PRE); Levyns 1282 Acocks 2312 (PRE 2 sheets, S); 3 km south of Piquetberg (BOL); along road from Clanwilliam to Algeria (–BB), (–DD), Ihlenfeldt 1023 (PRE); 2 mls [3.22 km] along Boatwright et al. 207 (JRAU); Olifants Dam (–BB), Piquetberg to Velddrif road (–DD), Marsh 795 (NBG, Compton 20015 (BOL, NBG 2 sheets); Clanwilliam PRE). 3219 (Wuppertal): pass into Biedouw Valley, south of town next to Olifants River (–BB), De Vos 6.2 mls [9.98 km] from turnoff from Clanwilliam– 1717 (NBG, PRE); near Clanwilliam on road to Bie- Calvinia road (–AA), Marsh 388 (K, NBG, PRE); bot- douw Valley (–BB), Joffe 959 (PRE); Ramskop Wild tom of Biedouw Pass (–AA), Mauve & Oliver 89 Flower Garden (–BB), Le Roux 2476 (BOL, NBG); (NBG, PRE); Pakhuis (–AA), Salter 1660 (BM, BOL, rocky hills near Clanwilliam (–BB), Parker 3589 (BOL, K); Pakhuisberg (–AA), Schlechter 139 (PRE); Schlech K, NBG); Pakhuis Pass (–BB), Schutte 262, 264 (JRAU); ter 8657 (PRE); Schlechter 8659 (BM, K, S); Pakhuis Bakhoogte near Clanwilliam (–BB), Stirton 9195 Pass (–AA), Stokoe s.n. sub SAM 52520 (NBG); Pakhuis (PRE); Nardouwskloof Farm (–BB), Stirton 9364 Pass, east side near Big Bends and gravel pit (–AA), Tay (PRE); Boschkloof (–BB), Thorne s.n. (NBG); between lor 11560 (NBG, PRE); Uitkyk Pass, into Biedouw val- Koperfontein and Malmesbury (–BC), Barker 4067 ley (–AA), Thompson 2926 (K, NBG); Cederberg State (BOL, NBG); on Citrusdal road 8 mls [12.87 km] from Forest, Grootkloof (–AC), Andrag 272 (NBG, PRE); Clanwilliam (–BD), Gillett 4082 (BOL, PRE); Olifants Nieuwoudt Pass (–AC), Barnes s.n. (BOL); Olifants S T R E L I T Z I A 39 (2018) 45

River near Algeria (–AC), Boatwright et al. 119 (JRAU) (NBG); hillside Voordenberg, near De Wet station (– Cederberg Reserve (–AC), Hubbard 34 (NBG); Wup- CB), Walters 2481 (NBG); between Worcester and Stet- perthal, just outside town on road to Clanwilliam (– tyns (–CD), Rycraft 1732 (NBG); De Doorns (–CD), AC), Grobbelaar 2975 (PRE); Wupperthal (–AC), Stir Moss 6798 (BM, J), Schmidt s.n. (NBG); Stettyns (–CD), ton 6399 (K, PRE); Garskloof, near Algeria (–AC), Van Rensburg 386 (K, NBG 2 sheets, PRE); Farm Dor- Stirton 9192 (PRE); near turn-off to Algeria on old Na- ingkloof, southern foothills of Voetpadsberg (–DA), tional road (–AC), Van der Walt s.n. (NBG, PRE); ca. Morley 506 (NBG); Hex River Kloof (–DA), Sidey 1878 8 km from Algeria on road to Clanwilliam (–AC), Van (PRE, S 2 sheets); Farm Doringkloof, foothills of Voet- Wyk 2568 (JRAU 2 sheets, PRE); Cedarberg, Forest padsberg (–DA), Van Wyk 66 (NBG, PRE, UPS); Jo- Reserve, Langrug (–AC), Viviers 510 (NBG, PRE); naskop (–DC), Bayliss BRI1228 (K, PRE); on plains be- Brakfontein (–AD), Ecklon & Zeyher 1332 (K, PRE, S); tween Assegaaikloof and Breede River (–DC), Bowie s.n. south of Trekkloof Pass (–AD), Low 2942 (NBG); (BM); Robertson (–DD), Levyns 2812 (BOL); Vrolik- Wuppertal (–AD), Thode A1987 (K, PRE); Thorne s.n. heid Nature Reserve (–DD), Van der Merwe 2969a (K, (NBG); Van Rooyen et al. 696 (NBG); Warm Baths, PRE). 3320 (Montagu): Kogmanskloof (–CC), Court Clanwilliam division (–CA), Edwards 252 (PRE); near 399 (GRA), Gillett 4521 (BOL); Mundt s.n. (K); Bon- Citrusdal (–CA), Steyn 391 (NBG 2 sheets); just north- nievale (–CC), Du Preez 321 (NBG); Rothman east of Citrusdal (–CA), Strid & Strid 37340 (NBG). NBG221/34 (NBG); Van Breda & Joubert 1948 (K); near 3318 (Cape Town): Riebeek Wes (–BD), Coffer s.n. Hassaquaskloof and Kogmanskloof (–CC), Ecklon & (NBG); Riebeeck Castle (–BD), Drège s.n. ‘III, D, a’ Zeyher 1333 (K, S); 15 mls [24.14 km] beyond Montagu (BM, K, S); Pillans 9788 (BOL); Stark & Co. s.n. (BOL). on Barrydale road (–CC), Marais NBG525/62 (NBG 2 3319 (Worcester): Farm Welbedacht, Biedouw Vally (– sheets); Bonnievale, hills of Bokkeveld beds (–CC), Mar AA), Johannesburg Botanical Garden 3876 (PRE); Hex loth 11811 (NBG 2 sheets, PRE); 5 km before Montagu River (–BC), Bolus 5152 (K); Marloth 1605 (PRE); (–CC), Van Wyk 544 (NBG, PRE); Goudmyn, between MacOwen 1605 (BM, K, NBG, UPS); main road pass- Bonnievale and Robertson (–CC), Van Wyk 2705 ing De Doorns, east of town (–BC), Mauve & Oliver (JRAU); Olifantsberg (–CC), Viviers 422 (NBG 2 136 (NBG, PRE); Hex River Pass (–BD), Eliovson 318 sheets, PRE); Scheepersrus (–CD), Johnson 147 (NBG 2 (J); Gray s.n. sub BOL 26052 (BOL, K); 5 mls [8.05 km] sheets); Barrydale (–DC), Galpin 3922 (GRA, PRE), from Worcester on Bains Kloof road (–CA), Story 2902 Morris 161 (BOL, NBG 2 sheets); Wall 3 (S 2 sheets); (PRE); Fairy Glen Hiking trail, Worcester (–CB), Cupi 15 mls [24.14 km] west of Barrydale (–DC), Gillett 1897 do 146 (NBG); 3.5 mls [5.63 km] from Worcester on (BOL, K); 20 km from Barrydale on road to Montagu (– road to Rawsonville (–CB), Grobbelaar 1183 (PRE); DC), Grobbelaar 2246 (PRE); 47 km from Montagu on 6 km from Worcester on road to Ceres (–CB), Grobbe road to Barrydale (–DC), Grobbelaar 2782 (K, PRE); laar 2796 (K, PRE); Worcester Veld Reserve (–CB), shaly hills at Barrydale (–DC), Hafström & Acocks 2314 Henrici 3807 (BOL, NBG); Midgley & Bosenberg 7 (PRE); 19 mls [30.57 km] south of Barrydale (–DC), (NBG 2 sheets, PRE); Olivier 101 (NBG); Van Breda & Martin 403 (NBG 2 sheets); alongside road up to Wilde- Joubert 2177 (K); Farm Vrolikheid, McGregor (–CB), hondskloof Pass between Montagu and Barrydale (– Jooste 192 (NBG); Blinkberg Pass, Tuinkloof (–CB), DC), McDonald 1568 (NBG, PRE); crossroad at Barry- Stirton 5907 (K); National road bridge at Breede River dale (–DC), Nel s.n. sub STE 18105 (NBG); between (–CB), Taylor 3729 (BOL, K, NBG); Karoo Botanical Barrydale and Montagu (–DC), Rycraft 2627 (NBG 3 Garden, Worcester (–CB), Barker 5944 (BOL, NBG 3 sheets); Van Niekerk 582 (K); top of Wildehondskloof sheets); Dobay 52 (NBG); Niemand 14 (JRAU); Van Pass (–DC), Van Wyk 2651b (JRAU). 3321 (Ladi- Wyk 1408a (JRAU); Fairy Glen, lower slopes of Brand- smith): Gamka Mountain Reserve, alongside road to wag Mountains (–CB), Stirton 9143 (PRE); Brandwagt, Paardebond (–CB), Cattell & Cattell 169 (K, NBG, in Fairy Glen Kloof (–CB), Van Breda SKF574 (K, PRE). 3322 (Oudtshoorn): 7 mls [11.26 km] south-west PRE); Robertson veld (–CB), Walters 380 (NBG); of De Rust (–BC), Theron 2071 (BOL, K, PRE, UPS); roadside Worcester west en route Brandwacht (–CB), Oudtshoorn (–CA), Besselaar s.n. (NBG); along roadside Walters 588 (NBG); Brandwacht (–CB), Walters 1237 from Oudtshoorn to De Rust (–CA), Boatwright et al. 46 S T R E L I T Z I A 39 (2018)

114 (JRAU); Farm Die Krans (–CB), Dahlstrand 2004 (K); 8 km from Stormsvlei to Swellendam (–AA), Stir (NBG, PRE); Dahlstrand 2312 (NBG, PRE); Oudts ton 6147 (K, PRE); Stormsvlei (–AA), Taylor 4048 hoorn, near Farm Oude Muragie (–CB), Vlok 2150 (NBG 2 sheets); Swellendam (–AB), Burchell 7475 (K); (PRE); Farm Doornkraal, 3 mls [4.83 km] east of De barren mountain ridges on the Riviersonderend at Rust (–DA), Dahlstrand 1469 (J, PRE). 3418 (Simons Storm Valley, Hassaquaskloof and Breede River (–AB), town): near Steenberg’s Cove, Vredenburg (–BB), Tay Zeyher 2320 (PRE 3 sheets). 3421 (Riversdale): near lor 1535 (NBG 2 sheets). 3419 (Caledon): Brakfontein Platte Kloof (–AA), Muir 671 (PRE); Roadside near (–BB), Schlechter 5279 (BM, BOL, S 2 sheets, PRE); Riversdale (–AB), Bayliss BRIB422 (PRE); Martin Krommerivier (–BB), Shaw 5624 (BOL, K); Rivierson- 9649 (GRA). Precise locality unknown: Cape, Bowie derend Mountains at Stormvallei, Assegaaikloof and s.n. (K); without locality, Drège s.n. (K, PRE); e Cap. b. Breede River (–BB), Zeyher 2320 (K, PRE 2 sheets). Spei, Sparrman s.n. (S); Thunberg s.n. (S); Thunberg s.n. 3420 (Bredasdorp): Breede River (–AA), Mundt 37 sub UPS 16413, 16414 (UPS). S T R E L I T Z I A 39 (2018) 47

6. Calobota elongata (Thunb.) Boatwr. & B.-E.van Wyk, S. African J. Bot. 75: 553. 2009. Basionym: Crotalaria elongata Thunb., Prodr. Pl. Cap.: 125. 1800. TYPE: SOUTH AF- RICA, ‘Carro prope Bockeveld’ [grid unknown], Thunberg s.n. sub THUNB-UPS 16544 (lectotype, designated here, UPS!). Lebeckia melilotoides R.Dahlgren, Bot. Not. 120: 268–271. 1967. TYPE: SOUTH AF- RICA, Platfontein, east of Hottentots Holland Kloof [3319 BB], H. Hall 177 (holotype, NBG!; isotypes, NBG!, PRE!, S!).

Erect to procumbent, virgate, multi-stemmed, 6–7 mm long. Pods ovate to elliptic, terete, unarmed shrub up to 1 m in height. Branches densely pubescent, subsessile, 5–8 × 1.5– green or light brown, young and older branch- 4.0 mm, 1- to 2-seeded, indehiscent; endocarp es densely sericeous. Leaves digitately trifoliol- glabrous. Seeds reniform, 2.4–3.0 × 1.3– ate, persistent; petiole longer or shorter than 2.0 mm, mature seeds light brown to light leaflets, 8–29 mm long; leaflets widely obo- pink, often mottled with brown or grey, sur- vate, alternate, sericeous, subsessile, terminal face smooth (Figure 15). Flowering time: late leaflet 13–32 × 8–20 mm, lateral leaflets 10– spring to summer (October to November). 32 × 6–16 mm, apex obtuse or retuse, base cuneate. Inflorescence 80–432 mm long, spicate, Distribution and habitat with more than 100 flowers; flowers sessile, This species occurs between Touws River bract 5–7 mm long, linear, densely tomen- and Ceres on sandy soil or dunes at ca. 600 m tose; bracteoles 2–5 mm long, linear, densely and has been recorded to be both avoided and tomentose. Flowers 8–11 mm long, pale yel- grazed by livestock (Figure 16). low. Calyx 4.5–6.0 mm long, tomentose; tube 3.5–5.0 mm long; lobes 1.1–1.7 mm long, deltoid. Standard 10–12 mm long; claw IUCN Red List category linear, 2.5–3.5 mm long; lamina narrowly The conservation status of this species was as- ovate to elliptic, 7.5–9.0 × 5–6 mm, pilose sessed as Vulnerable (VU D1+2) by Raimondo along dorsal midrib; apex obtuse. Wings et al. (2009). Calobota elongata is largely re- 10.5–12.0 mm long, claw 3.5–4.0 mm long, stricted to the Tanqua Karoo and it is estimat- lamina oblong, as long as or shorter than keel, ed that less than 1 000 individuals are left. It is 6–8 × 2.5–3.5 mm, glabrous, with 5–6 rows threatened by overgrazing and building activi- of sculpturing. Keel 9.5–12.0 mm long, ties. claw 3.0–4.5 mm long, lamina boat-shaped, 6.5–8.0 × 3.5–4.5 mm, glabrous. Pistil subsessile to shortly stipitate, densely pubescent, Discussion ovary linear to elliptic, 5.5–6.5 × 0.6–1.0 mm This species is superficially similar to Calo with 3 to 6 ovules; style ± as long as the ovary, bota obovata, but differs in the habit (a virgate 48 S T R E L I T Z I A 39 (2018)

A B1 B2

G H

I J1 J2 J3 K1

Figure 15.—Morphology of Calobota elongata. A, leaf in abaxial view; B1, bracts; B2, bracteoles; C, flower in lateral view; D, outer surface of the calyx (upper lobes to the left); E, standard petal; F, wing petal; G, keel petal; H, androecium; I, pistil; J1, long, basifixed anther; J2, intermediate carinal anther; J3, short, dorsifixed anther; K1, pod in lateral view; K2, pod in dorsal view. A–B, D–J:Van Wyk 2229, JRAU; C: Hall 177, NBG; K: Van Breda 4486, PRE. Scale bars: A & K, 10 mm; B–J, 1 mm. S T R E L I T Z I A 39 (2018) 49

the ovary and the pods are velvety and many- seeded). The growth form of C. elongata is very interesting and deserves further study. Field observations showed that new flowering stems are formed at ground level around the periphery of the plant each year and these remain quite prostrate until anthesis. As fruit formation progresses, the branches curl upwards and inwards, so that the small fruits are dispersed from some distance above the ground. The central part of the plant comprises infertile and senescent stems from previous seasons.

Figure 16.—Distribution of Calobota elongata. Thunberg s.n. sub THUNB-UPS 16544 is the only specimen in Thunberg’s Herbarium; it is annotated on the reverse side of the specimen by him and so is here designated as lecto- shrub with unbranced flowering stems aris- type of Calobota elongata. ing at ground level), trifoliolate leaves, dense, spicate inflorescences with smaller, sessile flowers, long, narrow bracts and bracteoles, Additional specimens very short calyx lobes, ovate to elliptic stand- examined ard petal with a pilose midrib, wing petals SOUTH AFRICA. 3219 (Wuppertal): Ceres Karoo, that are longer than keel, fewer ovules in the near Beukes Fountain (–DC), Schlieben & Van Breda ovary and shorter, tomentose pods with less 9811 (K, NBG, PRE). 3319 (Worcester): Verkeerde seeds (in C. obovata, a much-branched woody vlei (–BD), Van Wyk 2229 (2 sheets), 2230, 2562b (2 shrub, the leaves are unifoliolate, the inflores- sheets), 2710 (JRAU); Farm Littouw (–CB), Van Bre cences racemose, the flowers pedicellate, the da 4461 (PRE). 3320 (Montagu): near Railway line ca. bracts and bracteoles ovate, the calyx lobes 5 km south of Touws River (–AC), Van Breda 4486 (PRE); Leopard Rock Private Nature Reserve, Doorn- longer, the standard petal densely pubescent rivier (–DD), Vlok 2842 (NBG). Precise locality un- and widely ovate and the wing petals shorter known: south end of Tanqua Karroo, Story 3606 (K, than the keel, numerous ovules present in PRE). 50 S T R E L I T Z I A 39 (2018)

7. Calobota halenbergensis (Merxm. & A.Schreib.) Boatwr. & B.-E.van Wyk, S. African J. Bot. 75: 553. 2009. Lebeckia halenbergensis Merxm. & A.Schreib., Bull. Jard. Bot. État Bruxelles 27: 276. 1957. TYPE: NAMIBIA, Lüderitz South, Halenberg [2615 CB], Dinter 6632 (holotype, M, photo!; isotypes, BM! 2 sheets, K!, NBG!, PRE!, S!, Z, photo!). Lebeckia decutiens var. β glabrata E.Mey., Comm. Pl. Afr. Austral. [Meyer] 1: 34. Feb. 1836. TYPE: SOUTH AFRICA, ‘Karroo inter Goedemanskraal et Kaus’ [2917 AD], altit. 2000 ped., Drège s.n. ‘III, B’ (lectotype, designated here, P!).

Multi-stemmed shrub up to 1.5 m in height. shortly stipitate, glabrous; ovary linear, 4.5– Branches green; young branches pilose; older 7.5 × 0.8–1.1 mm with (8–)13 to 18(–22) branches pilose or glabrous with light brown ovules; style shorter than ovary, 2–4 mm bark. Leaves digitately trifoliolate, persistent; long. Pods linear, terete, glabrous, subsessile petiole 18–40 mm long, longer than leaflets; to shortly stipitate, 16–32 × 2–4 mm, ±3- to leaflets narrowly oblanceolate to linear, alter- 7-seeded, dehiscent; endocarp hairy. Seeds ob- nate, pilose on both surfaces, subsessile, termi- long reniform to reniform, sometimes some- nal leaflet 5–30 × 0.5–1.5(–2.0) mm, lateral what angular, 2.0–3.2 × 1.3–2.0 mm, mature leaflets 4–16 × 0.5–1.0 mm, apex acute, base seeds light brown to light orange, often mot- cuneate. Inflorescence 47–155 mm long, with tled with brown, surface smooth (Figure 17). 10 to 20 flowers; pedicel 1–2 mm long; bract Flowering time: specimens in flower and fruit 1–3 mm long, linear, pubescent on outer sur- have been collected from March to October; face; bracteoles 0.5–1.5 mm long, linear, pu- the main flowering period appears to be late bescent on outer surface. Flowers 7–11 mm winter to spring (June to October). long, yellow. Calyx (3.5–)4.0–6.0 mm long, pilose to glabrescent on outer surface; tube Distribution and habitat 2.5–4.0 mm long; lobes 1–2 mm long, sub ulate. Standard 7.0–11.5 mm long, claw Calobota halenbergensis occurs from the Lü- linear, 2–4 mm long, lamina widely ovate to deritz area in southwestern Namibia south- orbicular, 4–7 × 4.5–8.0 mm, pilose along wards into South Africa as far as Garies, south dorsal midrib; apex obtuse. Wings 8–11 mm of Springbok (Figure 18). It can be found on long, claw 2.5–4.0 mm long, lamina oblong, well-drained sandy loam, rocky sandstone, longer than keel, (4.5–)5.5–7.0 × (2.2–) gravelly or loose red sand often in dry river- 2.5–3.5 mm, glabrous, with 6–9 rows of beds or along disturbed roadsides at altitudes sculpturing. Keel 7.0–9.5 mm long, claw of between ca. 50 and 1 000 m above sea level. 2.5–4.0 mm long, lamina boat-shaped, 4.0– According to Mucina et al. (2006) it is an im- 5.5 × 2.5–4.0 mm, glabrous or sometimes portant component of Namaqualand Harde pilose on terminal parts. Pistil subsessile to veld (SKn 4 Namaqualand Heuweltjieveld) S T R E L I T Z I A 39 (2018) 51

A B C

D E F

G H1 I H2

K1 J1 J2 J3

Figure 17.—Morphology of Calobota halenbergensis. A, leaf in abaxial view; B, outer surface of the calyx (upper lobes to the left); C, flower in lateral view; D, standard petal; E, wing petal; F, keel petal; G, pistil; H1, bract; H2, bracteoles; I, androecium; J1, long, basifixed anther; J2, intermediate carinal anther; J3, short, dorsifixed anther; K1, pod in lateral view; K2, pod in dorsal view. A, C–F, H–K2:Boatwright et al. 146, JRAU; B: Middlemost 2129, NBG; G: Salter 5549, PRE. Scale bars: A & K, 10 mm; B–J, 1 mm. 52 S T R E L I T Z I A 39 (2018) and Namaqualand Sandveld (SKs 7 Nama- qualand Strandveld, SKs 8 Namaqualand Coastal Duneveld, SKs 11 Namaqualand Arid Grassland, SKs 12 Namaqualand Spi- nescent Grassland). Specimen label information indicates that it is grazed by livestock.

IUCN Red List category Considered of Least Concern (LC) by Rai- mondo et al. (2009). The conservation status of Calobota halenbergensis in Namibia has been assessed as lower risk (LR-nt) by Gold- ing (2002). Figure 18.—Distribution of Calobota halenber gensis. Discussion Calobota halenbergensis is similar to C. an gustifolia, but differs in the pilose leaves, gla- Bay (–AB), Powrie LWP1163 (WIND); Klinghardt brous keel petals and glabrous, fewer seeded Mountains, dark rock mountain east of a patch of shift- ing dunes in the valley ca. 5 km northeast of Sargdeckel, pods (and pistil). In C. angustifolia the leaves on the road south to Oranjemund from Lüderitz in the are sericeous, the keel petals pilose and the Diamond Area 1 (–BC), Bean & Oliver 2445 (BOL, pods (and pistil) sericeous with more seeds. NBG, PRE); Klinghardt Mountains (–BC), Dinter 3987 (BOL); Müller 830 (PRE, WIND); Namit- The specimen Drège s.n. ‘III, B’ in P displays sas, south of Klinghardt basin (–BD), Mannheimer the type locality and the characteristic gla- CM2107 (WIND); 20 km east of Kakaoberg Escarp- ment (–DB), Jürgens 28153 (PRE); Boegoeberg (–DD), brous pods of Calobota halenbergensis and is Dinter 6552 (BM). 2716 (Witputz): Klinghardt Moun- therefore chosen as lectotype of Lebeckia de tains, koppie southwest face (–AC), Mannheimer cutiens var. b glabrata. CM1455 (WIND); Diamond area 1, Polly Mountain (–AD), Wendt 20/2 (WIND); Udabib Mountain (– BB), Müller 810 (K, PRE, WIND); sandy flats at the Additional specimens base of Rooiberg sandstone mountain (–CB), Bean & examined Oliver 2386 (NBG, PRE); Diamond Area 1, Aurusberg (–CB), Burke 96225 (WIND); Oliver 10112 (WIND); NAMIBIA. 2615 (Lüderitz): Halenberg (–CB), Dinter Sperrgebiet, central Aurus Valley (–CB), Jürgens 28205 3805 (BM, BOL 2 sheets, K, NBG); 23 km from Rot- (PRE); Aurus at new camp (–CB), Mannheimer kop Station on power line track (–DC), Kolberg & CM2418 (WIND); Farm Spitzkop (–DC), Giess 14639 Maggs 184 (WIND). 2616 (Aus): plains around Garub (WIND); Wiss 2059 (WIND 3 sheets); Zebrafontein (–CA), Strohbach 577 (WIND). 2618 (Keetmans (–DD), Merxmüller & Giess 3310, 28861 (K, PRE, hoop): river at Spitzkop (–AD), Wiss 2059 (WIND). WIND); Müller & Horn 1615 (WIND); Rosh Pinah, 2715 (Bogenfels): 9.8 km south of Grillental on main west of airfield (–DD), Zietsman 1831 (WIND). 2816 north–south road between Lüderitz and Alexander (Oranjemund) 15 km west of Rosh Pinah on road to S T R E L I T Z I A 39 (2018) 53

Obib (–BA), Giess 13820 (K, WIND); Diamond Area Goldblatt 6002 (K, PRE, S); Van Wyk 3086 (JRAU 1, Valley between Obib Mountain Peak and Obib dunes 5 sheets); Van Wyk 6189 (PRE); 10 mls [16.09 km] (–BA), Van Wyk 9034 (PRE, WIND); 40 km from northwest of Steinkopf (–BC), Salter 5549 (BM, BOL, Brandkaros (–BC), Petersen s.n. (NBG). K, PRE); 5 mls [8.05 km] south of Komaggas (–CA), Compton 22794 (NBG); Sandkop gate at Kleinsee SOUTH AFRICA. 2816 (Oranjemund): between (–CA), Le Roux 5183 (NBG); Komaggas (–CA), Brandkaros and Grootderm (–BC), Venter 8199 (PRE); Maguire 400 (NBG 4 sheets); Grootmis, at Springbok Numees (–BD), Jürgens 22149 (PRE); McDonald turnoff on Port Nolloth–Kleinsee road (–CA), Rösch 739 (NBG, PRE); Annisvlakte (–BD), Jürgens 22322 & Le Roux 560 (PRE); just northeast of Grootmis (– (PRE); Oranjemund (–CB), Merxmüller & Giess 2266 CA), Strid & Strid 37734 (NBG); between Spektakel (BM, K, PRE, WIND); 70 km north of Oranjemund and Kleinsee (–CB), Grobbelaar 1962 (PRE); Farm (–CB), Williamson 4900 (WIND); 38 mls [61.14 km] north of Port Nolloth, on way to Alexander Bay (–DA), Langhoogte on Port Nolloth turn-off from Springbok– Werger 519 (K); Goariep-Passage (–DB), Jürgens 22485 Kleinsee road just north of waterpipe (–CB), Le Roux (PRE); Goariepvlakte (–DB), Jürgens 23119 (PRE); 3959 (NBG); Spektakel (–DA), Compton 11508 (NBG, near Doringbank, between Lekkersing and Kuboes (– PRE); Esterhuysen s.n. sub PRE 53706 (PRE); bot- DB), Van Wyk 2836 (JRAU, 4 sheets); Holgat River tom of Wildepaardehoek Pass (–DC), Van Jaarsveld (–DB), Wisura 2499 (NBG); Sukkel (–DD), Jürgens 1392 (PRE). 2918 (Gamoep): Karuchab [Arragaab], 22739 (PRE); between Port Nolloth and Holgat (– Richtersveld (–BB), Herre s.n. sub STE 11887 (NBG); DD), Pillans 5191 (K). 2817 (Vioolsdrif): bed of Ku- Karuchabpoort, Lekkersing (–BB), Oliver, Tölken & bus River (–AC), Hardy & Venter 4757 (PRE, WIND); Venter 145 (PRE). 3017 (Hondeklipbaai): Farm Ko- northwest slope of Noemiesberg down to Grasvlakte (– ingnaas 475 on road from Hondeklipbaai to Koingnaas AD), Smook 7937 (PRE); in flats between Blackhills and (–AB), Le Roux & Lloyd 353 (NBG); ca. 6 km north Wildepaardenrant (–CA), Thompson & Le Roux 288 of Koingnaas on Springbok road (–AB), Van Wyk 2848 (K, NBG); Eksteensfontein (–CA), Venter 8053 (PRE); (JRAU, 2 sheets); 3 km southwest of Soebatsfontein on 5 mls [8.05 km] west of Stinkfontein (–CC), Leist road to Wallekraal (–BA), Le Roux 2899 (NBG); Farm ner 3400 (K, WIND); Lekkersing (–CC), Thorne s.n. Doornfontein 464, Portion 1, Steenkamp Kraal (–BA), (NBG); 7 mls [11.26 km] beyond Stinkfontein on way Le Roux 4557, 4630 (NBG); on road from Garies to to Kuboos, near Skouerfontein (–CC), Werger 431 (K, Hondeklipbaai (–BC), Boatwright et al. 146 (JRAU); PRE, WIND); Blomhoek, 5 km from Eksteensfontein Wallekraal, between Garies and Hondeklipbaai (–BC), on road to Kubus (–DD), Germishuizen 4723 (PRE). Compton 5485 (BOL, NBG); Wallekraal (–BC), Pil 2916 (Port Nolloth): 16 km north of Port Nolloth lans 17970 (BOL); 5 km from Wallekraal to Garies (– on road to Alexander Bay (–BB), Germishuizen 5315 BC), Stirton 6053 (K); Horees, ca. 30 mls southwest (BOL, PRE); 38 mls [61.14 km] east by south of Port of Kamieskroon (–BC), Van Breda 4085 (PRE); Farm Nolloth (–BD), Acocks 14236 (K, PRE 2 sheets); Port Sandkraal, 40 km west of Garies (–DA), Hilton-Taylor Nolloth (–BD), Bolus 427 (BM, BOL, K, NBG, PRE, 1371 (NBG, PRE); 12.9 km from Garies–Hondeklip- UPS); 2 mls [3.22 km] east of Port Nolloth (–BD), baai road on road to Soutfontein, Farm Sandkraal (– Marloth 12661 (NBG, PRE); between Port Nolloth DA), Steiner 2172 (NBG, PRE); in red sand near Garies road and Stinkfontein (–BD), Middlemost 2129 (NBG (–DB), Esterhuysen s.n. (K, NBG 2 sheets, PRE). Pre- 2 sheets, S); 25 mls [40.23 km] after Anenous Pass (– cise locality unknown: between Port Nolloth and Au- BD), Schlechter sub STE 10890 (NBG 2 sheets, K, PRE). grabies Poort, Bolus 6457 (K); Doornpoort, Compton 2917 (Springbok): 37.5 km from Port Nolloth on road 20633 (NBG 3 sheets); Kalkfontein, near river, Marloth to Steinkopf (–AA), Clarke 640 (K, PRE); Anenous 12273 (NBG 4 sheets, PRE); kloof by Doornpoort, Flats along road to Eksteenfontein (–AB), Goldblatt & Richtersveld, Herre s.n. sub STE 11865, 11866 (NBG); Manning 9284 (NBG); on road to Port Nolloth from Noagas, Drège s.n. ‘III, B’ (BM, K, P); sandveld, Rich- Steinkopf (–AC), Boatwright et al. 149, 190 (JRAU); tersveld, Herre s.n. sub STE 11868 (NBG 2 sheets, PRE); Anenous Pass (–BA), Germishuizen 4838 (PRE); without locality, Drège s.n. sub PRE 9367 (PRE). 54 S T R E L I T Z I A 39 (2018)

8. Calobota linearifolia (E.Mey.) Boatwr. & B.-E.van Wyk, S. African J. Bot. 75: 553. 2009. Basionym: Lebeckia linearifolia E.Mey., Comm. Pl. Afr. Austral. [Meyer] 1: 33. Feb. 1836. TYPE: SOUTH AFRICA, Northern Cape Province, on the Gariep, near Verleptpraam [2817 AA], 300 ped., Drège s.n. ‘III, B’ (lectotype, designated here, P!; isolectotypes, K!, P!, S!, Z photo!). Lebeckia dinteri Harms, Repert. Spec. Nov. Regni Veg. 16: 360. 1920. TYPE: NAMIBIA, Garub [2616 CA], Dinter 1057 (lectotype, designated here, NBG!).

Erect or virgate, multi-stemmed, unarmed somewhat clavate, terete, sericeous, subsessile shrub up to 3 m in height. Branches green; to shortly stipitate, 25–40 × 2–3 mm, ±4- to young branches sericeous or silky; older 8-seeded, dehiscent; endocarp hairy. Seeds branches sericeous or silky with light brown oblong-reniform, 2.5–3.5 × 1.5–2.5 mm, bark. Leaves simple, persistent; petiole ab- mature seeds light brown or grey, surface sent; leaflets linear to oblanceolate or spathu- smooth (Figure 19). Flowering time: mainly late, sericeous, sessile, leaflets 11–45 × 1.0– between September and November, but 2.5 mm; apex acute; base cuneate. Inflorescence flowering and fruiting specimens have been 38–170 mm long, with 5 to 9 flowers; pedicel collected in January, April, May and August. 1–2(–8) mm long; bract 1.5–2.5 mm long, linear to narrowly ovate, pubescent; brac- Distribution and habitat teoles 1.0–1.5 mm long, linear to narrowly ovate, pubescent. Flowers 13–17 mm long, Calobota linearifolia is widely distributed in bright yellow. Calyx 5.5–7.5 mm long, seri- the Northern Cape Province of South Africa ceous; tube 4–6 mm long; lobes 1.0–2.5 mm and southern Namibia, with outlier popula- long, deltoid. Standard 11.5–15.0 mm long; tions around the Unjab River, Sima Hill and claw linear, 2.5–3.5 mm long; lamina ovate, the Sanitatas area in northwestern Namibia 8.5–12.0 × 8.5–10.5 mm, dorsal surface pu- (Figure 20). Polhill (2003) mentioned that bescent; apex acute. Wings 8.0–11.5 mm this species also occurs in Botswana in the long, claw 3.0–4.5 mm long, lamina oblong, Kgalagadi district, but no specimens from shorter than keel, 5–7 × 2.5–4.0 mm, pi- this region were found in the herbarium col- lose, with 4–8 rows of sculpturing. Keel 12– lections studied. Calobota linearifolia occurs 15 mm long, claw 3.5–6.0 mm long, lamina in dune sand, red sand or brackish soil often boat-shaped, 7.5–10.0 × 4.0–5.5 mm, pu- along rivers or in dry watercourses, washes bescent. Pistil subsessile to shortly stipitate, and gulleys. It occurs at altitudes of between pubescent; ovary linear, 9–11 × 0.8–1.5 mm ca. 300 and 1 350 m. It is suspected of poi- with 12 to 18 ovules; style shorter than ovary, soning livestock (according to specimen label 4.8–7.8 mm long. Pods linear, sometimes information). S T R E L I T Z I A 39 (2018) 55

A1 A2 B C

D E F

G H1 H2 I

J1 J2 J3 K1

Figure 19.—Morphology of Calobota linearifolia. A1, A2, leaves in abaxial view; B, flower in lateral view; C, standard petal; D, outer surface of the calyx (upper lobes to the left); E, wing petal; F, keel petal; G, pistil; H1, bract; H2, bracteoles; I, androecium; J1, long, basifixed anther; J2, intermediate carinal anther; J3, short, dorsifixed anther; K1, pod in lateral view; K2, pod in dorsal view. A1, B: Acocks 13195, PRE; A2: Giess & Van Vuuren 821, BOL; C: Giess 13803, WIND; D–G, J: Dean 673, JRAU; H, I: Evrard 9285, PRE; K: Keet 1668, WIND. Scale bars: A & K, 10 mm; B–J, 1 mm. 56 S T R E L I T Z I A 39 (2018)

IUCN Red List category Considered of Least Concern (LC) by Rai- mondo et al. (2009).

Discussion Calobota linearifolia is similar to C. cinerea, but differs in its virgate to erect habit, simple, linear leaves, fewer flowered inflorescences, sericeous pods that are constricted between the seeds and the light brown to grey seeds. Calobota cinerea has a low, spreading habit, obovate, elliptic or oblanceolate and trifoliolate leaves and the inflorescences more densely Figure 20.—Distribution of Calobota linearifo flowered. The latter species also has tomen- lia. tose pods that are not constricted between the seeds and the seeds are light brown to orange and often mottled. (–AC), Tinley 1625 (WIND). 2013 (Unjab Mouth): 8 mls [12.87 km] east of Torrabay (–AA), Giess et al. The P specimen of Drège s.n. ‘III, B’ bears 6180 (PRE, WIND); Unjab River (–AB), Kers 1875 the type locality and was annotated by Meyer (S). 2517 (Gibeon): Orion, Varsfontein (–AC), Giess et al. 7190 (K); between Achterfontein and Gelwater himself and is therefore chosen as lectotype (–BC), Pearson 9233 (K). 2520 (Mata Mata): Kala- of Calobota linearifolia. No other specimens hari Gemsbok National Park (–CA), Schlieben 10433 except the one in NBG could be traced of the (PRE). 2615 (Luderitz): Farm Gawachab KEE 114 (– collection Dinter 1057 (it is not present in ei- BB), De Winter & Giess 6424 (BOL, K, WIND). 2616 ther B or Z). Both Dinter 1056 and Dinter (Aus): between Neisip and Eureka (–BC), Merxmül 1058 were found in Z, but it is likely that all ler & Giess 2986 (WIND); 13 mls [20.92 km] west of Aus (–CA), Giess & Van Vuuren 821 (BOL, K, specimens of Dinter 1057, except the NBG WIND); Kuibis (–DA), Bayliss BRI.B.365 (K, PRE specimen chosen as lectotype, have been 2 sheets). 2617 (Bethanie): Felshang, near Seeheim destroyed. (–DD), Merxmüller & Giess 3615 (PRE, WIND); Seeheim (–DD), Dinter 1205 (NBG). 2618 (Keet- manshoop): 7 mls [11.26 km] south by east of Naute Additional specimens Station (–DB), Acocks 15659 (K, PRE); Sandmodder examined 73 (–DD), Lensing A87/75 (WIND). 2619 (Aroab): east of Groot Karas Mountains (–CA), Mannheimer NAMIBIA. 1812 (Sanitatas): 20 mls [32.18 km] et al. CM771 (WIND). 2715 (Bogenfels): Sperrgebiet south of Orupembe (–CD), Giess & Leippert 7435 South, 38 km east of Kakaoberg escarpment (–DB), (WIND). 1913 (Sesfontein): 2 km north of Sima Hill, Jürgens 28176 (PRE); Chameis Valley (–DC), Wil Kaokoland (–AA), Braine 68 (WIND); near Sima Hill, liamson 2647 (BOL). 2717 (Chamaites): Mara, Let- northeast of Möwe Bay (–AA), Moss & Jacobsen K316 terklip area in Konkiep River bed (–CD), Craven 2634 (PRE, WIND); between Hoamib and Unjab Rivers (WIND); bed of Vis River, near Seeheim (–DA), Keet S T R E L I T Z I A 39 (2018) 57

1663 (WIND, 2 sheets). 2719 (Tränental): Warmfon- sheets); Farm Aluriesfontein 308 (–DA), Owen-Smith tein 280 (–AB), Lensing J14/76 (WIND); Müller 87 1252 (WIND). 2819 (Ariamsvlei): Nababis, Guigatsis (WIND); Farm Blaufontein WAR 276 (–AC), Giess et (–BA), Evrard 9220 (PRE). 2820 (Kakamas): below al. 7131 (WIND). Riemvastmaak, Upington (–AD), Barnard s.n. (NBG). SOUTH AFRICA. 2620 (Twee Rivieren): Ashkam 2821 (Upington): Swartpan, northwest of Upington (–DD), Evrard 9285 (PRE). 2720 (Noenieput): (–AA), Snijman 214 (NBG); 24 mls [38.62 km] south- 20 mls [32.18 km] north of Noenieput on way to east of Groblershoop (–DD), Leistner & Joynt 2861 Kalahari Gemsbok Park (–AD), Werger 1489 (NBG, (K, PRE). 2822 (Glen Lyon): 9 mls [14.48 km] west PRE); Askam area, 120 km from Van Zylsrus (–BB), south-west of Volop (–CD), Leistner 1742 (K). 2824 Dlamini et al. EvW568 (K). 2723 (Kuruman): 10 mls (Kimberley): Gordonia (–BA), Webb 16 (K, PRE). [16.09 km] southwest of confluence of the Kuruman 2918 (Gamoep): Karasberg (–CB), Van Schauroff and Molopo Rivers (–AA), Leistner 2255 (K). 2817 s.n. (K). 2921 (Kenhardt): 40 mls [64.36 km] north (Vioolsdrif): 5 km west of Nuob River (–AA), Giess northeast of Kenhardt (–AC), Acocks 18803 (K). 3020 13803 (K, PRE, WIND); Garip (–AA), Wyley s.n. (S); (Brandvlei): sand dunes along Sak River near Brandvlei Richtersveld (–BA), Venter 8069 (PRE). 2818 (Warm- (–AD), Acocks 13195 (BOL, K, PRE). 3022 (Carnarv- bad): a little south of Dabegabis (–BC); Pearson 4390 on): Van Wyksvlei, Beesdam (–AD), Dean 673 (JRAU (BM 2 sheets, K); near Dabegabis (–BC), Pearson 4896 2 sheets). Precise locality unknown: without local- (K); mountains between Wolftoon and Henkriesfon- ity, Bolus 1935 (BOL), Flanagan s.n. sub PRE 53667 tein (–CC), Pearson 3105 (BM 2 sheets, BOL, K 2 (PRE), Range 1935 (BOL, NBG). 58 S T R E L I T Z I A 39 (2018)

9. Calobota lotononoides (Schltr.) Boatwr. & B.-E.van Wyk, S. African J. Bot. 75: 553. 2009. Basionym: Lebeckia lotononoides Schltr., Bot. Jahrb. Syst. 27: 143. 1900, as ‘L. lotonoides’; Germishuizen & Meyer, Plants of Southern Africa: an annotated checklist, Strelitzia 14: 523. 2003, p.p., Boatwr. & B.-E. van Wyk, in S. African J. Bot. 73 (4): 664–666. 2007, as ‘L. lotononoides’, non Goldblatt & Manning. 2000. TYPE: SOUTH AFRICA, ‘In regione namaquensi: In sabulosis montium Karree-Bergen, alt. C. 4500 ped.’ [3118 AB], Schlechter 8214 (lectotype, designated here, K!; isolectotypes, BM!, BOL!, MO, photo!, PRE!, S!, Z, photo!).

Small, decumbent, multi-stemmed, unarmed 5.5–7.0 × 3.0–4.5 mm, glabrous. Pistil sub- shrublet up to 0.5 m in height. Branches sessile to shortly stipitate, pubescent, ovary green; young branches sericeous, older linear, 6.5–7.5 × 0.7–0.9 mm with 15 to 19 branches pilose to glabrous with light brown ovules; style shorter than ovary, 3.0–3.5 mm bark, often buried. Leaves digitately trifoliol- long. Pods narrowly oblong, sometimes some- ate or rarely 5-digitate with two additional what clavate, sericeous, terete, subsessile to leaflets, persistent; petiole 10–23 mm long, shortly stipitate, 18–37(–40) × 3–4 mm, 7- to longer than leaflets; leaflets elliptic to nar- 12-seeded, dehiscent; endocarp hairy. Seeds rowly oblanceolate, secund, alternate, seri- suborbicular, 2.0–2.5 × 1.5–2.0 mm, mature ceous, conduplicate, subsessile, terminal seeds brown with beige spots, hilum brown, leaflet 10–17 × 1.5–3.0 mm, lateral leaflets surface rugose (Figure 21). Flowering time: 8–14 × 1.5–3.0 mm, mucronate, base nar- July to October, late winter to late spring. rowly cuneate. Inflorescence (15–)24–54 mm long, with (3–)4 to 14 flowers; pedicel Distribution and habitat 1–3 mm long; bract 0.7–1.5 mm long, linear, pubescent; bracteoles 0.3–0.6 mm long, Calobota lotononoides occurs on the West linear, pubescent. Flowers 12–15 mm long, Coast of South Africa around Hondeklipbaai yellow. Calyx 5.0–6.5 mm long, pubescent; and Brand-se-baai (Figure 22). The type collec- tube 3.0–3.5 mm long; lobes 2–3 mm long, tion was made by Schlechter at Karree-Bergen, subulate. Standard 13–16 mm long; claw close to Nuwerus in the Vanrhynsdorp district linear, 4–7 mm long; lamina widely ovate to according to Leistner and Morris (1976). The transversely oblong, 8–9 × 9.0–12.5 mm, pi- species occurs on well-drained, sandy soils with lose along dorsal midrib; emarginate. Wings the main stems becoming covered with sand 14–15 mm long; claw 3.0–4.5 mm long; lam- (Boatwright & Van Wyk 2007). The multi- ina narrowly oblong, subfalcate, twice as long stemmed, decumbent habit appears to be an as the keel, 11.5–15.0 × 2–4 mm, glabrous, adaptation to moving sand dunes, as extensive with 7–8 rows of sculpturing. Keel 9–11 mm underground branches enable the plant to long; claw 3–4 mm long; lamina boat-shaped, emerge above the sand after being covered. S T R E L I T Z I A 39 (2018) 59

B A1 A2 C

D E

G1 G2 H I

J1 J2 J3

Figure 21.—Morphology of Calobota lotononoides. A1, A2, leaves in abaxial view; B, flower in lateral view; C, standard petal; D, wing petal; E, keel petal; F, pistil; G1, bract; G2, bracteoles; H, outer surface of the calyx (upper lobes to the left); I, androecium; J1, long, basifixed anther; J2, intermediate carinal anther; J3, short, dorsifixed anther; K1, pod in lateral view; K2, pod in dorsal view. A–K: Boatwright et al. 142, JRAU. Scale bars: A & K, 10 mm; B–J, 1 mm. 60 S T R E L I T Z I A 39 (2018)

IUCN Red List category According to Raimondo et al. (2009) the status of this species is Near Threatened (NT B1ab(iii)). Although only a 5% habitat loss has been observed due to heavy mineral sand mining, the loss is still ongoing and could later become problematic.

Discussion Calobota lotononoides superficially resembles C. angustifolia, notably the broad-leaved form on the West Coast and around Clanwil- liam. In this form the leaflets are broad and Figure 22.—Distribution of C. lotononoides. sericeous, much like those of C. lotononoides. However, C. lotononoides differs in its smaller, decumbent habit with buried stems and especially in the exceptionally long lamina of the Additional specimens wing petals, which is ± twice as long as those examined of the glabrous keel petals. The inflorescences SOUTH AFRICA. 3017 (Hondeklipbaai): from are unarmed, and the sericeous pods contain Sandkraal, north of Bitter River, 25 km west of Garies spotted, rugose seeds. Calobota angustifolia (–CB), Helme et al. 4720 (NBG); Farm Roodeheuw- can reach heights of up to 2 m and the wing el, 9 km west of Nariep (–DC), Boatwright et al. 142 petals are only slightly longer than the pilose (JRAU); Perold 1646 (PRE). 3117 (Lepelfontein): keel petals. Vredendal, southeast of Brand-se-baai, on Farm Harte- beestekom (–BD), Helme 2931 (NBG); Vanrhynsdorp/ Vredendal, Brand-se-baai (–BD), Van Rooyen 2235 A large number of Schlechter’s African col- (PRE). 3118 (Vanrhynsdorp): Karree-Bergen (–AB), lections are housed at K (Stafleu & Cowan Schlechter s.n. sub TRV 1059 (PRE); from Skaapvlei, 1985) and we therefore select the K specimen some 10 km northeast of Olifants River mouth (–AC), as lectotype of Calobota lotononoides. Helme 4684 (NBG). S T R E L I T Z I A 39 (2018) 61

10. Calobota namibensis Boatwr. & B.-E.van Wyk, sp. nov. TYPE: NAMIBIA, 25 mls [40.23 km] east of Lüderitz on road to Aus [2615 CA], De Winter & Hardy 7919 (holotype, WIND!, specimen on the right; isotypes, K!, PRE!). Lebeckia dinteri auct. non Harms, Repert. Spec. Nov. Regni Veg. 16: 360. 1920.: A.Schreib., FSWA 60: 67. 1970.

Virgate, multi-stemmed shrubs up to 1.2 m flowering specimens have been collected in tall. Branches green; young branches seri- March, August and October. ceous; older branches sericeous or glabrous with light brown bark. Leaves simple, de- Distribution and habitat ciduous; petiole absent; elliptic to ovate, upper and lower surfaces sericeous, alter- Calobota namibensis is endemic to Namibia nate, 6–12 × 1–2 mm; apex acute; base cu- and occurs in the Lüderitz and Aus areas neate. Inflorescences 70–110 mm long, with (Figure 24). It grows on dunes in red sand 3 to 8 flowers; pedicel 1–2 mm long; bract or gravelly sandy soil along roadside up to 2.5–3.5 mm long, linear, sericeous; brac- 1 800 m above sea-level. teoles 1.5–3.0 mm long, linear, sericeous. Flowers 10–11 mm long, bright yellow. IUCN Red List category Calyx 6–8 mm long, densely pubescent on the outer surface; tube 3–4 mm long; lobes The status of this species has been assessed as 3.0–4.5 mm long, subulate. Standard 9.5– Vulnerable (VU D2) by Golding (2002; in- 12.0 mm long; claw linear, 2–3 mm long; cluded as Lebeckia dinteri) and as Least Con- lamina ovate, 7.5–9.0 × 5–6 mm, densely cern (LC) recently by Loots (2005; included pubescent on dorsal surface; apex acute. as Lebeckia dinteri). Wings 8.0–8.5 mm long; claw 2–3 mm long; lamina oblong, shorter than the keel, Discussion 5.5–6.5 × 2.5–3.0 mm, with 6–8 rows sculpturing, glabrous. Keel 8.0–11.5 mm long; Calobota namibensis is a very poorly known claw 2.5–3.5 mm long; lamina boat-shaped, and collected species. It is similar to C. lineari 5.5–8.0 × 3.5–4.5 mm, pubescent. Pistil folia and C. saharae, but differs in the elliptic to shortly stipitate, pubescent; ovary linear, ovate leaves, densely pubescent petals and the 4–5 × 0.9–1.0 mm with about 5 ovules; style short, tomentose pods. In both C. linearifolia longer than the ovary, 5.0–9.5 mm long. Pods and C. saharae the leaves are linear. The pet- linear, terete, 15–17 × 2 mm, 3- to 4-seeded, als are also pubescent in C. linearifolia while tomentose, ?dehiscent, endocarp glabrous. in C. saharae only the standard petal is pilose. Seeds not seen (Figure 23). Flowering time: The pods of C. linearifolia are sericeous, while 62 S T R E L I T Z I A 39 (2018)

A B D

D E F

J1 J2 J3

H1 H2 I

Figure 23.—Morphology of Calobota namibensis. A, flower in lateral view; B, standard petal; C, outer surface of the calyx (upper lobes to the left); D, leaf in abaxial view; E, wing petal; F, keel petal; G, pistil; H1, bract; H2, bracteoles; I, androecium; J1, long, basifixed anther; J2, intermediate carinal anther; J3, short, dorsifixed anther; K1, pod in lateral view; K2, pod in dorsal view. A–C, E, F, H, J: Owen-Smith 1268, WIND; D: Kolberg & Maggs 212, WIND; G, I: Owen-Smith 117, WIND; K: De Winter & Hardy 7919, WIND. Scale bars: A & K, 10 mm; B–J, 1 mm. S T R E L I T Z I A 39 (2018) 63 those of C. saharae are glabrous and in both species they are longer than those of C. na mibensis.

Harms (1920) described Lebeckia dinteri from a specimen that is actually Calobota linearifo lia. Subsequent authors, however, referred to the species here described as new as Lebeckia dinteri. The species is quite distinct from Calo bota linearifolia, but is here formally described for the first time.

Additional specimens examined Figure 24.—Distribution of Calobota namiben sis. NAMIBIA. 2615 (Lüderitz): dunes south of Koichab pump station (–AD), Seely 2005 (WIND); Koichab State Land (–BC), Owen-Smith 117, 1268 (WIND); Halenberg (–CB), Merxmüller & Giess 3088 (WIND); Rotkop, Lüderitz (–CB), Dinter 1032 (NBG); Vuuren 821 (PRE, WIND); koppie southwest of Rot- Merxmüller & Giess 3141 (WIND). 2616 (Aus): kop Station, next to power-line track (–CB), Kolberg & 13 mls [20.92 km] west of Aus (–CB), Giess & Van Maggs 212 (PRE, WIND). 64 S T R E L I T Z I A 39 (2018)

11. Calobota obovata (Schinz) Boatwr. & B.-E.van Wyk, S. African J. Bot. 75: 553. 2009. Basionym: Lebeckia obovata Schinz, Mem. Herb. Boissier 1: 126. 1900. TYPE: NAMIBIA, Gansberg [grid unknown], Fleck 75 (lectotype, Z-000022914, photo!, specimen on the left, designated here; isolectotype, Z-000022915, photo!).

Sericeous, multi-stemmed shrub up to 1.5 m specimens have been collected in May, June, in height. Branches green; young branches July, November and December suggesting velvety, older branches velvety or glabrous that flowering occurs in spring and summer. with light brown bark. Leaves unifoliolate or rarely trifoliolate, persistent; petiole shorter Distribution and habitat than leaflets, 5–24 mm long; leaflets widely obovate to orbicular, rarely ovate, alternate, Calobota obovata is endemic to Namibia and velvety, subsessile, 12–36 × 10–25 mm, apex occurs from Windhoek southwards as far as obtuse or rarely acute, base cuneate. Inflores Aus (Figure 26). It grows on rocky outcrops cence 24–85 mm long, with 7 to 16 flowers; or granite at altitudes of ca. 1 600 to 2 230 m. pedicel 4–5 mm long; bract 3–5 mm long, ovate, pubescent; bracteoles 3–5 mm long, IUCN Red List category ovate to linear, pubescent. Flowers 13–16 mm long, bright yellow. Calyx 8.0–8.5 mm long, The status of this species has been assessed as densely pubescent; tube 4.5–5.5 mm long; Data Deficient (DD) by Golding (2002) and lobes 3–4 mm long, subulate. Standard Loots (2005). 14.5–15.5 mm long, claw linear, 3.5–4.5 mm long, lamina widely ovate, 10.0–11.5 × 12.5– Discussion 14.5 mm, dorsal surface pubescent; apex obtuse. Wings 14.5–15.5 mm long, claw The species is superficially similar toCalobota 3.0–3.5 mm long, lamina oblong, shorter elongata, but differs in the unifoliolate leaves, than keel, 11–12 × 5.5–6.0 mm, pilose, with few-flowered racemose inflorescences, pedi- 4–5 rows of sculpturing. Keel 14.5–16.5 mm cellate flowers, obovate bracts and bracteoles, long, claw 4–5 mm long, lamina boat-shaped, widely ovate standard petal, pubescent wing 10.0–11.5 × 5.5–6.5 mm, pubescent. Pis petals that are shorter than the keel petals and til subsessile to shortly stipitate, pubescent, the linear, sericeous pods (C. elongata differs ovary linear, 11.5–12.5 × 1.5–2.0 mm with in the trifoliolate leaves, dense, spicate inflo- ±15 ovules; style shorter than ovary, 6–7 mm rescences with much smaller, sessile flowers, long. Pods linear, terete, subsessile to shortly long, narrow bracts and bracteoles, ovate to stipitate, 30–40 × 4–5 mm, 5- to 7-seeded, elliptic standard petal with a pilose midrib, velvety, dehiscent; endocarp hairy. Seeds not wing petals that are longer than keel and seen (Figure 25). Flowering time: flowering shorter, tomentose pods with less seeds). The S T R E L I T Z I A 39 (2018) 65

A1 A2 C

D E F

J 1 J2 J3

G1 G2 H

K 1 I

K 2

Figure 25.—Morphology of Calobota obovata. A1, A2, leaves in abaxial view; B, flower in lateral view; C, standard petal; D, wing petal; E, keel petal; F, outer surface of the calyx (upper lobes to the left); G1, bract; G2, bracteoles; H, androecium; I, pistil; J1, long, basifixed anther; J2, intermediate carinal anther; J3, short, dorsifixed anther; K1, pod in lateral view; K2, pod in dorsal view. A1: Schweidfezer 4315, WIND; A2, K: Dinter 1094, NBG; B, F, I: Pearson 8036, BOL; C–E, G, H, J: Kers 152, WIND. Scale bars: A & K, 10 mm; B–J, 1 mm. 66 S T R E L I T Z I A 39 (2018) growth forms of the two species are also very different. Calobota obovata is an erect, much- branched, shrub, while C. elongata has a virgate habit, with slender, mostly unbranched flowering stems arising from a woody base at ground level.

According to Stafleu and Cowan (1985), Hans Schinz’s main collection is in Z and we therefore designate one of the Z specimens as lectotype.

Additional specimens examined Figure 26.—Distribution of Calobota obovata. NAMIBIA. 2217 (Windhoek): Farm Regenstein (– CA), Giess & Giess 11765 (NBG, PRE, S, WIND); Merxmüller & Giess 28010 (WIND); Ruschberg (–CA), Rusch 4379 (S). 2316 (Nauchas): Road to 2318 (Leonardville): Farm Rietfontein (–CA), Leip Hakos, just off the C26 to Walvis Bay, Gamsberg (– pert 4537 (WIND). 2516 (Helmeringhausen): Farm AC), Boatwright et al. 241 (NBG, WIND); Gamsberg Aruab 23 on top edge of Great Escarpment (–DA), (–AD), Kers 152 (WIND); Kers 154 (S); Merxmül Miller Mil1/036 (WIND). 2615 (Lüderitz): Lüderitz ler & Giess 957 (BM, K, PRE, WIND); Kolberg (–CA), Rogers 2959 (K). 2616 (Aus): Urus Plateau, & Tholkes NAM2960-HK1418 (K); Meyer 1016 Lüderitz (–BA), Logan & Seely 301 (WIND); Aus (– (WIND); Schweidfezer 4315 (WIND); Farm Hope- CB), Dinter 1094 (NBG); Aus, rocks east of railway field (–AD), Merxmüller & Giess 893 (WIND); Farm station (–CB), Pearson 8036 (BM, BOL 2 sheets, K, Weipenfels (–AD), Walter & Walter 1686 (WIND). NBG, PRE). S T R E L I T Z I A 39 (2018) 67

12. Calobota psiloloba (E.Mey.) Boatwr. & B.-E.van Wyk, S. African J. Bot. 75: 554. 2009. Ba- sionym: Stiza psiloloba E.Mey. Comm. Pl. Afr. Austral. [Meyer] 1: 32. Feb. 1836. Lebeckia psiloloba (E.Mey.) Walp., Linnaea 13: 478. 1839. TYPE: SOUTH AFRICA, s. loc., Drège 6470 (lectotype, designated here, P!; isolectotype, K!, S!). Acanthobotrya pungens sensu Eckl. & Zeyh., Enum. 2: 193. Jan 1836 [non Lebeckia pun gens Thunb., Nov. Gen. Pl. [Thunberg]: 141. 1800]. TYPE: SOUTH AFRICA, ‘Zwartkops– et Zondagsrivier’, Ecklon & Zeyher 1340 (lectotype, designated here, NBG!; isolectotypes, P!, K!, S!).

Erect, multi-stemmed, spinescent shrub up glabrous. Pistil stipitate, glabrous; ovary linear to 1.2 m tall. Branches green; young branch- to elliptic, 5.5–7.5 × 0.8–1.1 mm; ovules 6 to es glabrous to very sparsely pubescent, older 8; style shorter than ovary, 3.5–5.0 mm long. branches glabrous with brown bark. Leaves Pods oblong, laterally compressed, membra- unifoliolate or rarely trifoliolate, decidu- neous, 14–17(–33) × 4–5 mm, glabrous; ous; petiole short, 0.5–1.0 mm long (up to stipe up to 4 mm long, 2- to 6-seeded, inde- 5 mm long when trifoliolate); leaflets slightly hiscent; endocarp glabrous. Seeds reniform, obovate to spathulate, alternate, upper and 2.5–3.0 × 2.0–2.5 mm, light brown, surface lower surfaces thinly pilose, 5–7(–11) × 1.5– smooth (Figure 27). Flowering time: the main 3.0(–4.0) mm; base attenuate; apex obtuse. flowering time is between March and June, Inflorescences 40–50(–65) mm long, with 8 but flowering also sometimes occurs in spring to 10(–12) flowers; pedicel relatively long, (3– and summer (October to January). )4–5(–6) mm long; bract 0.3–0.4(–0.6) mm long, triangular; bracteoles 0.1–0.3 mm long, Distribution and habitat linear to slightly ovate. Flowers 8–11 mm long; bright yellow. Calyx 2.2–2.6 mm long, Calobota psiloloba is restricted to the Eastern slightly pubescent on the outer surface; tube Cape Province and occurs mainly around Port 1.5–2.0 mm long; lobes widely triangu- Elizabeth, Grahamstown and Addo, but has lar, 0.5–0.8 mm long; apices broadly acute. outlier populations around Cradock and Mid- Standard 7–9 mm long; claw 2.5–3.0(– delburg at altitudes between 30 and 520 m 4.0) mm long; lamina oblong to slightly ovate, (Figure 28). It is normally found in open grass- 4–5(–6) × 3.0–4.5 mm, glabrous; apex emar- lands on sandy soils and is eaten by elephants ginate. Wings 7.0–8.5 mm long; claw 2.5– (according to specimen label information). 3.0(–3.5) mm long; lamina oblong, shorter than the keel, 4.0–4.5(–5.0) × 2–3 mm, with IUCN Red List category 4–5 rows sculpturing, glabrous. Keel 8.5– 11.0 mm long; claw 3.5–4.0(–4.5) mm long; Considered to be of Least Concern (LC) by lamina boat-shaped, 5–6(–7) × 3–4 mm, Raimondo et al. (2009). 68 S T R E L I T Z I A 39 (2018)

A B

G1 G2 J1 J2 J3 I

K1 H1 H2

Figure 27.—Morphology of Calobota psiloloba. A, flower in lateral view; B, standard petal; C, outer surface of the calyx (upper lobes to the left); D, wing petal; E, keel petal; F, pistil; G1, G2, leaves in abaxial view; H1, bract; H2, bracteoles; I, androecium; J1, long, basifixed anther; J2, intermediate carinal anther; J3, short, dorsifixed anther; K1, pod in lateral view; K2, pod in dorsal view. A: Story 3353, PRE; B–F, G1, H–J: Van Wyk s.n., JRAU; G2: Zeyher 2317, NBG; K: Le Roux et al. 20, JRAU. Scale bars: G & K, 10 mm; A–F, I, J, 1 mm. S T R E L I T Z I A 39 (2018) 69

of this species. The specimen of Ecklon & Zeyher 1340 in NBG dislays the type locality as the Enumeratio label and is therefore chosen as lectotype of Acanthobotrya pungens.

Additional specimens examined SOUTH AFRICA. 3125 (Steynsburg): Grootfontein, Middelburg district (–AC), Burtt-Davy 9665 (PRE). 3225 (Somerset East): Cradock (–BA), Botany Hon ours Students 320 (PRE, PRU); Somerset East, farm Ka- boega (–DA), Bayliss 6897 (K, S). 3324 (Steytlerville): on plains near Zondags River (–DB), Bowie s.n. (BM). Figure 28.—Distribution of Calobota psiloloba. 3325 (Port Elizabeth): in the valleys of the Zuurberg Mountains at Paarde Poort (–AC), Bolus 2667 (BOL, K); Wilson 422 (BM); Perdepoort (–AC), Theron 685 (PRE); Bassons Kloof, Annes Villa (–BB), Bayliss 3923 (NBG); Addo Elephant Park (–BD), Archibald 3848 Discussion (GRA, K, PRE); Brynard 445 (K, PRE); Repton 5715 th Calobota psiloloba is similar to C. cuspidosa, (PRE); 15 mile [24.1 km] on the road from Port Eliza- but differs in that this species only reaches beth to Addo (–BD), Long 612 (GRA, K); along road from Grahamstown to Paterson (–BD), Van Wyk s.n. heights of up to 1.2 m and has small flowers (JRAU); Uitenhage (–CD), Alexander-Prior s.n. (BM, that are borne on long (3–6 mm long), thin K 3 sheets); Cradock Place, Port Elizabeth (–DC), Gal pedicels (C. cuspidosa reaches heights of up pin 6430 (GRA, PRE); district Uitenhage, Zwartkops to 4 m and has much larger flowers on rela- River (–DC), Ecklon & Zeyher 737 (BM, BOL 2 sheets, tively short pedicels). The older stems are al- K 2 sheets, NBG, PRE 4 sheets); Ecklon & Zeyher 1340 most completely glabrous and the standard (K, NBG, P 3 sheets, PRE, S); New Brighton near Port Elizabeth (–DC), Ethel West 395 (BOL); Coega (–DC), petal notched, while in C. cuspidosa the older Long 945 (GRA, K); Redhouse, near cemetary (–DC), stems are pubescent and the standard petal is Troughton 425 (PRE 2 sheets); Redhouse (–DC), Boat obtuse. The pods and seeds are also smaller wright et al. 107 (JRAU); Le Roux et al. 20 (JRAU); than those of C. cuspidosa. Wells 3697 (PRE); Zwartkops River (–DC), Zeyher 2317 (PRE 2 sheets, NBG, S). 3326 (Grahamstown): The specimen of Drège 6470 at P bears a label Table farm near Grahamstown (–BC), Atherstone 24, with the words ‘Stiza psiloloba mihi’ in Mey- 50 (K); Grahamstown (–BC), MacOwen 1102 (K); Bushman’s River bridge, main road from Grahams er’s handwriting, and is therefore chosen as town to Port Elizabeth (–CA), Archibald 3958 (PRE); lectotype of C. psiloloba. The relatively thin, Story 3353 (K, PRE). Precise locality unknown: glabrous twigs and small, leathery pods (in an without locality, Barber s.n. (K); Bowie s.n. (K); Zeyher envelope on the specimen) are characteristic 3317 (PRE). 70 S T R E L I T Z I A 39 (2018)

13. Calobota pungens (Thunb.) Boatwr. & B.-E.van Wyk, S. African J. Bot. 75: 554. 2009. Basionym: Lebeckia pungens Thunb., Nov. Gen. Pl. [Thunberg]: 141. 1800. TYPE: SOUTH AFRICA, Western Cape Province, near Olifant’s River [close to Oudtshoorn] at Cannaland [now Little Karoo], Thunberg s.n. sub THUNB-UPS 16417 (lectotype, designated here, UPS!). Stiza erioloba E.Mey., Comm. Pl. Afr. Austral. [Meyer] 1: 31. Feb. 1836. TYPE: SOUTH AFRICA, Western Cape Province, near Klaarstroom [close to Prince Albert, 3322BC], near the foot of the ‘Groote Zwartebergen’ [Great Swartberg], Drège s.n. (lectotype, designated here, P!, specimen on the right; isolectotypes, K!, P!, S!).

Erect, multi-stemmed woody spinescent Pistil subsessile to shortly stipitate, pubescent; shrub up to 2.5 m tall. Branches green; young ovary linear, 6.0–8.5 × 0.8–1.0 mm; ovules 6 branches densely sericeous; older branches to 9; style shorter than ovary, 4.0–5.5 mm long. sericeous with brown bark. Leaves unifoliol- Pods oblong to slightly falcate, laterally com- ate or rarely trifoliolate, deciduous; petiole pressed, membraneous, 11–21(–28) × (2.5– short, 0.5–2.0 mm long (up to 5 mm long )3.0–4.0(–6.0) mm, densely tomentose; when leaves are trifoliolate); leaflets obovate to stipe very short (less than 1 mm long), ±4- to broadly obovate, upper and lower surfaces seri- 6-seeded, indehiscent; endocarp glabrous. ceous, alternate, 3–13(–18) × 2–5(–8) mm; Seeds reniform, 2.0–2.5 × 1.5–2.0 mm, surface base attenuate; apex retuse. Inflorescences (22– smooth (Figure 29). Flowering time: flowering )25–40(–55) mm long, with (4–)6–10 flow- occurs from spring to summer (September to ers; pedicel 2–4 mm long; bract 0.6–1.1 mm December). long, triangular; bracteoles 0.4–0.6 mm long, narrowly ovate. Flowers 10–13(–14) mm long, Distribution and habitat bright yellow. Calyx 2.5–3.7 mm long, pubescent on the outer surface; lobes triangular, Calobota pungens occurs from Laingsburg 0.4–1.0 mm long; tube 2–3 mm long, apices through Ladysmith and Oudtshoorn to Ba- acute. Standard 9.5–12.0 mm long; claw viaanskloof and Willowmore in the east at 3–4 mm long; lamina widely ovate, 6–7(– altitudes of between 790 and 1 100 m (Fig- 8) × 4–6(–9) mm, pilose along the dorsal mid- ure 30). It is typically a fynbos species and oc- rib; apex broadly acute. Wings 7–12 mm long; curs on rocky, loamy or quartzitic soils. claw (2.0–)3.5–5.0 mm long; lamina oblong, shorter than the keel, 4–6(–7) × 2–3(–4) mm IUCN Red List category with 4–5 rows of sculpturing, pilose. Keel 10– 14 mm long; claw 4.0–5.5 mm long; lamina Considered to be of Least Concern (LC) by oblong, 6.0–8.5 × (3.0–)4.0–5.5 mm, pilose. Raimondo et al. (2009). S T R E L I T Z I A 39 (2018) 71

A B C

D1 D2

E F

G I J1 J2 J3

H1 H2 K1

Figure 29.—Morphology of Calobota pungens. A, flower in lateral view; B, standard petal; C, outer surface of the calyx (upper lobes to the left); D1, D2, leaves in abaxial view; E, wing petal; F, keel petal; G, pistil; H1, bract; H2; bracteoles; I, androecium; J1, long, basifixed anther; J2, intermediate carinal anther; J3, short, dorsifixed anther; K1, pod in lateral view; K2, pod in dorsal view. A, B, D1, E–G, H2, I, K: Boatwright et al. 106, JRAU; C: Taylor 9386, NBG; D2: Barker 9948, NBG; H1: Acocks 15547, PRE; J: Van Wyk 3252, JRAU. Scale bars: D & K, 10 mm; A–C, E–J, 1 mm. 72 S T R E L I T Z I A 39 (2018)

Discussion Calobota pungens is similar to C. cuspidosa and C. psiloloba, but differs from both in the sericeous stems, sericeous obovate leaves, pilose petals, pubescent ovary and tomentose pods (in C. cuspidosa the stems are pubescent and the leaves linear to slightly obovate, while in C. psiloloba the stems are glabrous or slightly pilose and the leaves slightly obovate to spathulate. In both these species the ovary and pods are glabrous).

The specimen Thunberg s.n. sub THUNB- UPS 16417 is chosen as lectotype of C. pun Figure 30.—Distribution of Calobota pungens. gens as it is the only specimen in Thunberg’s Herbarium and the type locality is written on the back of the sheets in Thunberg’s PRE); Markotter 9951 (NBG); Congo Valley close to handwriting. The Drège specimen of Stiza Oudtshoorn (–AC), Dreyer 18618 (NBG); northern erioloba in P is chosen as lectotype because it foothills of Swartberg above farm Rietfontein (–AC), Vlok 1152 (K, PRE); Venterskloof near Klaarstroom displays the type locality as well as the char- (–AD), Taylor 9386 (K, NBG); 20 mls [32.2 km] west acteristic tomentose pods from which Meyer of Klaarstroom (–AD), Acocks 15547 (K, PRE); on denominated this species. road between Prins Albert and Klaarstroom (–AD), Van Wyk EvW0151 (K); Meiringspoort (–BC), Boat wright et al. 106 (JRAU); Levyns 6619 (BOL); Stokoe Additional specimens 61389 (NBG); outside Meiringspoort, past Klaar- examined stroom turn-off along roadsides (–BC), Boatwright et al. 111 (JRAU); foot of Swartberg Mountains, near SOUTH AFRICA. 3320 (Montagu): Cabidu (–AB), Klaarstroom (–BC), Drège s.n. (S, K); Klaarstroom, Bond 12112 (BOL, NBG); Matjiesfontein (–BA), Prince Albert (–BC), Barker 9948 (NBG 2 sheets). Compton 2714, 3315 (BOL, K); Compton 22233 (NBG 3323 (Willowmore): Hot Springs (–AC), Fourcade 2 sheets); Foley 58 (PRE); Marloth 10764 (NBG 2 5841 (NBG); Fourcade 6086 (BOL, NBG); flats be- sheets, PRE); top of Koppie near Matjiesfontein (– tween Hot Springs and Toorwater (–AC), Thompson BA), Thoday & Delf (NBG); Rooikloof (–BD), Van 1398 (PRE); lower slopes of Zuurberg at Georgida Wyk 2147 (JRAU). 3321 (Ladysmith): Farm Withoek (–AD), Fourcade 4410 (BOL, K); 7 mls [11.26 km] (–AC), Van Wyk 2154 (JRAU); road from Seweweeks south-west of Willowmore (–AD), Horn s.n. (PRE); Poort (–AD), Levyns 2334 (BOL); Touwsberg, farm 6–8 mls [9.6–12.8 km] south-east of Willowmore on Rietfontein (–CA), Germishuizen 6956 (PRE); Tou- Baviaanskloof road (–AD), Wells 2606 (PRE); 7 mls wsberg, farm Wolwefontein (–CA), Van Wyk et al. [11.2 km] on Baviaanspoort road from Willowmore (– 3447 (JRAU, PRE). 3322 (Oudtshoorn): 14 mls AD), Wells 2838 (GRA, PRE); 18 ml [28.8 km] from [22.5 km] east of Prince Albert (–AA), Sidey 1906 Uniondale on Willowmore road (–AD), Wells 3699 (PRE, S); Swartberg Pass (–AC), Bond 1537 (NBG, A & B (GRA, PRE); along road from Willowmore S T R E L I T Z I A 39 (2018) 73

to Baviaanskloof near Uniondale turnoff (–BC), Snij Uniondale (–CA), Fourcade 6255 (BOL); Piet-se- man 353 (NBG, PRE); Kammanassie hill sides 7 mls laagte on road to Uniondale (–CA), Van Wyk 3180 [11.3 km] from Uniondale (–CA), Fourcade 3594 (JRAU); dirt road between De Rust and Uniondale (BOL 2 sheets, K); Vet Vlei, 8 mls [12.8 km] from (–CA), Van Wyk 3252 (JRAU). 74 S T R E L I T Z I A 39 (2018)

14. Calobota saharae (Coss. & Durieu) Boatwr. & B.-E.van Wyk, S. African J. Bot. 75: 554. 2009. Genista saharae Coss. & Durieu in Coss. Bull. Soc. Bot. France 2: 247. 1855. Spar tidium saharae (Coss. & Durieu) Pomel, Nouv. Mat. Fl. Atl.: 173. 1874. TYPE: ALGE- RIA, D’Oran Province, Cosson s.n. (type P-COS, not seen).

Erect or virgate, multi-stemmed, unarmed Distribution and habitat shrub up to 2 m in height. Branches green; young branches densely pilose or glabrous; Calobota saharae is endemic to North Africa older branches pilose or glabrous with light and occurs in Morocco, Algeria, Libya and brown bark. Leaves simple, deciduous; petiole Tunisia. It is found on desert hills or dunes absent; linear, alternate, pilose on both surfac- at altitudes of ca. 200 m (Figure 32). The dis- es, sessile, 6–10 × 1–2 mm, apex acute, base junct distribution is remarkable, but a careful cuneate. Inflorescence 75–150 mm long, with consideration of morphological and molecu- 5 to 12 flowers; pedicel 1–2 mm long; bract lar evidence leaves no doubt that the species 1.5–2.0 mm long, linear, pubescent; brac- is best accommodated within the genus Calo teoles 0.5–1.0 mm long, linear, pubescent. bota (see Boatwright et al. 2009). Flowers 13–17 mm long, yellow. Calyx 4.5– 7.0 mm long, pilose; tube 3–4 mm long; lobes 1.5–3.0 mm long, subulate. Standard 13– IUCN Red List category 15 mm long, claw linear, 3.5–4.0 mm long, Unknown. lamina ovate, 9.5–11.0 × 7.5–8.0 mm, dorsal surface pubescent; apex obtuse. Wings 11.5– 14.0 mm long, claw 4.0–4.5 mm long, lamina Discussion oblong, longer than keel, 7–9 × 3–4 mm, Calobota saharae is similar to Calobota cus glabrous, with 4 rows of sculpturing. Keel pidosa and C. psiloloba in the laterally com- 12.5–13.0 mm long, claw 4–5 mm long, lam- pressed, membraneous pods, but differs in ina boat-shaped, 8.0–8.5 × 4.5–5.0 mm, gla- the unarmed habit, linear leaves and pu- brous. Pistil subsessile to shortly stipitate, glabrous; ovary linear, 9.0–11.5 × 1.0–1.3 mm bescent standard petal (in C. cuspidosa and with 7 to 8 ovules; style shorter than ovary, C. psiloloba the branches are strongly spines- 4.5–5.5 mm long. Pods oblong-linear, later- cent, the leaves slightly obovate to elliptic or ally compressed, membraneous, subsessile spathulate and the petals pubescent). Calo to shortly stipitate, 30–40 × 6–10 mm, 4- to bota saharae is also similar to C. namibensis, 6-seeded, indehiscent; endocarp glabrous. but differs in the linear leaves, the glabrous Seeds reniform to oblong-reniform, 2.5– keel petals and the membraneous pods (in 3.0 × 1.5–2.0 mm, mature seeds light brown, C. namibensis the leaves are elliptic to ovate, surface smooth (Figure 31). Flowering time: the keel petals densely pubescent and the flowering occurs from March to May. pods thick-walled and tomentose). S T R E L I T Z I A 39 (2018) 75

A B C

D E F

J1 J2 J3 G I

K1 H1 H2

Figure 31.—Morphology of Calobota saharae. A, flower in lateral view; B, standard petal; C, outer surface of the calyx (upper lobes to the left); D, leaf in abaxial view; E, wing petal; F, keel petal; G, pistil; H1, bract; H2, bracteoles; I, androecium; J1, long, basifixed anther; J2, intermediate carinal anther; J3, short, dorsifixed anther; K1, pod in lateral view; K2, pod in dorsal view. A, E, G: Keith 181, K; B, C, F, H–J: Hill s.n., K; D: Pitard 3276, K; K: Alleizette s.n. sub K 2007/014804, K. Scale bars: D & K, 10 mm; A–C, E–J, 1 mm. 76 S T R E L I T Z I A 39 (2018)

There are several Cosson specimens from various localities in the D’Oran Province in the K collection but it is unclear to us which of these are types.

Additional specimens examined NORTH AFRICA. Algeria: Aïn Sefra, Alleizette s.n. sub K 2007/014804 (K); Cosson s.n. sub K 000227004 (K); Hill s.n. (K); Bourgeau 210 (K 2 sheets); Le Cesve 5429 (BM); Rothschild & Hartert s.n. (BM); Samuels son 6994 (K); Constantine Province, Oued Souf, Cosson s.n. sub K 000227008 (BM, K); Djebel Milogh, Cosson s.n. sub K 000227003 (K); Oran Province, Aïn Sefifsifa, Cosson s.n. sub K 000227005 (K); Oran Province, Abi- od-Sisi-Cheihk, Cosson s.n. sub K 000227006 (K); Wadi Rummel, above Sidi Benour, Guichard KG/Lib/289 (K); El Ardja, Pitard 3275 (K); Oued el Khezoua/El Oued, Pitard 3276 (K). Tunisia: Nefta, Cosson s.n. sub K 000227007 (K). Libya: Aïn Zara, near Pripoli, Brown Figure 32.—Approximate distribution of Calo s.n. (K); Jebel Nefoussa, Azizia-Rhnem, Davis 49544 bota saharae. (K); Jebel, north of Jefren, Keith 181 (K). S T R E L I T Z I A 39 (2018) 77

15. Calobota sericea (Thunb.) Boatwr. & B.-E.van Wyk, S. African J. Bot. 75: 554. 2009. Ba- sionym: Lebeckia sericea Thunb., Nov. Gen. Pl. [Thunberg]: 143. 1800. TYPE: SOUTH AFRICA, ‘e Cap. b. Spei’ [grid unknown], Thunberg s.n. sub THUNB-UPS 16423 (lectotype, designated here, UPS!). Lebeckia flexuosa E.Mey., Comm. Pl. Afr. Austral. [Meyer] 1: 34. Feb. 1836; Walpers in Linnaea 13: 478. 1839. TYPE: SOUTH AFRICA, ‘In rupestribus prope Kwekri vier’ [grid unknown], altit. 700 ped., Drège s.n. ‘III, C’ (lectotype, designated here, P!; isolectotypes, K!, P!). Lebeckia decipiens E.Mey., Comm. Pl. Afr. Austral. [Meyer] 1: 35. Feb. 1836. TYPE: SOUTH AFRICA, Karroo prope Mierenkasteel [grid unknown], Drège s.n. ‘III, C’ (lectotype, designated here, P!; isolectotypes, BM!, K!, NBG!, P!, PRE!, S!).

Sericeous, multi-stemmed unarmed shrub sculpturing. Keel 12.5–19.5 mm long, claw up to 2 m in height. Branches green; young 3.5–7.0 mm long, lamina boat-shaped, 8.5– branches sericeous; older branches sericeous 13.0 × 4.5–6.5 mm, pilose on terminal parts. or glabrous with brown bark. Leaves digi- Pistil subsessile to shortly stipitate, sericeous; tately trifoliolate; persistent; petiole longer ovary linear, 8.5–15.0 × 0.8–1.5 mm with or shorter than leaflets, 13–55 mm long; 19 to 28 ovules; style shorter than ovary, leaflets elliptic to narrowly oblanceolate, al- 3.5–8.5 mm long. Pods linear, terete, subses- ternate, sericeous, subsessile, terminal leaflet sile to shortly stipitate, 30–50 × 3–4 mm, 7- (15–)17–50 × (1–)2–8 mm, lateral leaflets to 14-seeded, sericeous, dehiscent; endocarp 10–45 × (1–)2–5 mm, apex acute, base an- hairy. Seeds oblong-reniform, 2.3–3.5 × 1.7– gustate. Inflorescence 70–220 mm long, with 2.5 mm, mature seeds light brown to light 10 to 28 flowers; pedicel 2–4 mm long; bract pink, surface smooth (Figure 33). Flowering 2–7 mm long, linear, pubescent; bracteoles time: flowers mainly from May to October; 1–5 mm long, linear, pubescent. Flowers one specimen collected in March was in 13–24 mm long, pale or bright yellow. Ca flower. lyx 6.0–9.5 mm long, sericeous; tube 4.0– 6.5 mm long; lobes 1–4 mm long, subulate. Distribution and habitat Standard (11.5–)13.0–19.0 mm long; claw linear, 3–5 mm long; lamina widely ovate, Calobota sericea is widely distributed from 8.0–14.5 × 9.5–17.0 mm, pilose along dorsal Wupperthal in the south, as far north as the midrib; apex obtuse. Wings 12.0–17.5 mm Oranjemund area (Figure 34). It grows in long; claw 3.0–5.5 mm long; lamina oblong, well-drained sandy loam, granite, rocky sand, as long as or shorter than keel, 8–12 × 4.0– sandy dunes, in riverbeds or along disturbed 6.5 mm, glabrous, with (5–)8–14 rows of roadsides at altitudes of between 300 and 78 S T R E L I T Z I A 39 (2018)

B1 B2 A1 A2

E F

H G J1 J2 J3

I K1

Figure 33.—Morphology of Calobota sericea. A1, A2, leaves in abaxial view; B1, bract; B2, bracteoles; C, flower in lateral view; D, standard petal; E, wing petal; F, keel petal; G, outer surface of the calyx (upper lobes to the left); H, androecium; I, pistil; J1, long, basifixed anther; J2, intermediate carinal anther; J3, short, dorsifixed anther; K1, pod in lateral view; K2, pod in dorsal view. A1, B–J:Boatwright et al. 151, JRAU; A2, K: Van Wyk 3119, JRAU. Scale bars: A & K, 10 mm; B–J, 1 mm. S T R E L I T Z I A 39 (2018) 79

and pods (in C. cytisoides the apices of the leaflets are mucronulate, the pedicels longer and the flowers larger, the bracts and bracteoles are often ovate, the calyx glabrous and the pistil and pods glabrous).

Thunberg s.n. sub THUNB-UPS 16423 is here chosen as lectotype of Calobota sericea as it is the only specimen in Thunberg’s Her- barium. The specimen of Drège s.n. ‘III, C’ in P has the diagnostic features of Lebeckia flex uosa described by Meyer and a label with the type locality, so that it is here chosen as lectotype. The specimen of Drège s.n. ‘III, C’ at P Figure 34.—Distribution of Calobota sericea. is chosen as lectotype of Lebeckia decipiens as it bears the type locality and was annotated by Meyer himself. 1 200 m. This species, according to Mucina et al. (2006) and Jürgens (2006), is an im- Additional specimens portant component of Richtersveld (SKr 14 Southern Richtersveld Inselberg Shrubland), examined Namaqualand Hardeveld (SKn 1 Namaqua- SOUTH AFRICA. 2816 (Oranjemund): sand drifts land Klipkoppe Shrubland, SKn 2 Nama- on Witbank (–DC), Pillans 5542 (BOL); sandy coast qualand Shale Shrubland, SKn 6 Kamiesberg belt between Port Nolloth and Holgat River (–DD), Mountains Shrubland), Namaqualand Sand- Pillans 5627 (BOL, K). 2817 (Vioolsdrif): Kodaspiek (–AA), Oliver et al. 451 (K, NBG); Vandersterrberg veld (SKs 9 Namaqualand Inland Duneveld) northeast of Khubus north and southeast of the top of and Southern Namib Desert (Dn 1 Alexan- Helskloof and southeast of Paradysberg western slopes der Bay Coastal Duneveld). to summit (–AC), Oliver et al. 151 (NBG, K, PRE); Vioolsdrif, Ridge above Koeskop, summit of Vander sterberg, Richterveld National Park (–AC), William IUCN Red List category son & Williamson 5764 (NBG); Cornellsberg (–CA), Bean 1258 (BOL); Bean 1328 (S); Van Jaarsveld 11977 Considered to be of Least Concern (LC) by (PRE); Viviers 2046 (NBG); Jenkins Hill, 20 km from Raimondo et al. (2009). Eksteenfontein on road to Mount Stewart (–CB), Ger mishuizen 4679a (PRE 2 sheets); Spektakel Hill (– DA), Johnson 216 (NBG). 2916 (Port Nolloth): sandy Discussion plain 2 mls [3.22 km] east of Port Nolloth (–BD), Mar Calobota sericea is similar to C. cytisoides, but loth 12660 (NBG, PRE). 2917 (Springbok): along roadside on road from Port Nolloth to Steinkopf (– differs in the acute apices of the leaflets, short AC), Boatwright et al. 151 (JRAU); Klipfontein (– pedicels and smaller flowers, linear bracts and BA), Bolus 429 (BM, BOL, K, NBG, PRE), Bolus 6549 bracteoles, sericeous calyx and sericeous pistil (K); Hutchinson 904 (BM, BOL, K, PRE); Steinkopf 80 S T R E L I T Z I A 39 (2018)

(–BA), Herre s.n. sub STE 11889 (NBG); Meyer s.n. sub O'kiep (–DB), Scholtz 20 (NBG, PRE); Hester Malan STE9068 (NBG); Anenous Pass (–BA), Germishuizen Nature Reserve (–DB), Stirton 6004 (PRE); Struck 113 4839 (PRE, WIND); Schutte 591 (JRAU 2 sheets); Van (NBG); Van Wyk 5728 (PRE); 5 mls [8.05 km] north Wyk 6266 (PRE); Spitskop, 6 mls [9.65 km] south of of O’kiep (–DB), Van der Schijff & Schweickerd 5744 Steinkopf (–BD), Lewis 5487 (NBG 2 sheets); Klipfon- (K, PRE); 2 mls [3.22 km] north of Springbok (–DB), tein Koppie (–CA), Compton 5486 (BOL, NBG); Van Niekerk 240 (PRE, 2 sheets); 11 km from Spring- Stinkfontein Mountains southwest of Van Zylsrus (– bok to Pofadder (–DB), Van Wyk 3078 (JRAU 2 CA), Oliver et al. 625 (NBG, PRE); Spektakel (–DA), sheets); Modderfontein (–DB), Whitehead s.n. (S); Compton 11524 (NBG); Spektakel Pass (–DA), Eliovs 3 km east of Springbok (–DB), Zietsman & Zietsman on 37 (NBG); 25 km from Springbok towards Spek- 667 (PRE); Messelpad between Springbok and Walle- takelberg Pass (–DA), Glen 1463 (BOL, PRE); Spek- kraal (–DC), Compton 20664 (NBG 2 sheets); near Ko- takel Pass, 26 km west of Springbok (–DA), Goldblatt maggas (–DC), Eliovson 69 (J); north slopes of Rooi 3651 (PRE); top of Spektakelberg Pass (–DA), Grobbe berg above Buffels River on eastern approach to the laar 2584 (PRE); Farm Ezelfontein 214, 2 km north of Meselpad (–DC), Hilton-Taylor 2140 (NBG, PRE); ca. Naries farmstead (–DA), Le Roux 3233 (NBG); 11 km from Springbok on sand road to Hondeklipbaai 21.7 km from Springbok to Spektakel (–DA), Stirton (–DC), Joffe 96 (NBG, PRE); on road to Hondeklip- 6019 (K); 24 km from Springbok to Kleinsee (–DA), baai, 18 mls [28.96 km] from turn-off from national Stirton 10149 (NBG); 9 mls [14.81 km] southeast by road to Springbok (–DC), Van der Merwe 215 (NBG); east of Springbok (–DB), Acocks 19237 (K, PRE); 29 mls [46.66 km] from Springbok on road to Garies 8.5 mls [13.68 km] west by south of Springbok (–DB), (–DD), Grobbelaar 1119 (PRE); near Mesklip (–DD), Acocks 19550 (BOL, K, PRE); near O’kiep (–DB), Leighton 1156 (BOL); on roadside of N7 highway Barker 6244 (NBG 3 sheets); 1ml [1.61 km] south of 13 km south of Springbok, opposite 33.4 marker (– Springbok (–DB), Barker 8389 (NBG 2 sheets); 2 km DD), Powrie 652 (NBG 2 sheets, PRE); 30 mls east of Springbok (–DB), Bellamy 12 (PRE); Spring- [48.27 km] south of Springbok (–DD), Rynolds 5440 bok (–DB), Bester & Pretorius 8 (JRAU); Crosby 20 (K); 20 mls [32.18 km] southwest of Springbok (–DD), (PRE); Salter 895 (BM, K); Small et al. 82 (PRE); Van Verdoorn & Dyer 1791 (BOL, K 2 sheets). 2918 Blerk 7 (PRE); Van Hille 8 (BOL); Venter 8820 (PRE); (Gamoep): Zilverfontein (–CC), Drège s.n. (BM, K, P, Nababeep, road to O’kiep (–DB), Burrows 2950 S). 3017 (Hondeklipbaai): Diknek (–AD), Van Breda (GRA); ±8 km northeast of Nababeep on gravel road 4073 (PRE); Farm Doornfontein 464, Portion 1, Steen- to junction with tar road (–DB), Botha 3170 (PRE); kamp Kraal (–BA), Le Roux 4555 (NBG); on road to 2 mls [3.22 km] northeast of Springbok (–DB), Comp Leliefontein from Kamieskroon (–BB), Boatwright et ton 22012 (NBG); Concordia, near Springbok (–DB), al. 219 (JRAU); Grootvlei, hillsides, Kamieskroon (– Clarke 687 (PRE); Thorns s.n. (NBG); just outside BB), Compton 6839 (NBG); Skilpad Wild Flower Re- Nababeep on road to O’kiep (–DB), Germishuizen serve, 100m from house (–BB), Cruz 85 (NBG); Studer 4515 (PRE); Nababeep (–DB), Gess & Gess 89/90/67 Pass (–BB), Evrard 8960 (PRE); Kamieskroon (–BB), (GRA); Hester Malan Wild Flower Reserve north of Hutchinson 841 (BM, K 2 sheets, PRE); Salter 863 Oxaliskoppie (–DB), Le Roux 918, 1231 (PRE); Naba- (BM); Salter 865 (K); Thorne s.n. (NBG); Van Rooyen beep, near golf course (–DB), Le Roux 2674 (BOL); 2416 (PRE); portion of Wolwepoort 459, Skilpad 2 mls [3.22 km] north of Springbok (–DB), Lewis 3392 Farm, northwest of Kamieskroon (–BB), Johannesburg (NBG); Schelpe 240 (BM, BOL); 4 mls [6.44 km] west Botanical Garden 3877 (PRE); between Kamieskroon of Springbok (–DB), Macguire 369 (BOL, NBG 2 and Soebatfontein (–BB), Meyer 7331 (PRE); ± 4 km sheets); Springbok, weather station hill on west of town from Kamieskroon (–BB), Meyer 7349 (PRE); Kamies- (–DB), Manning 2774 (NBG); Okiep (–DB), Düm berg Pass (–BB), Schutte 593 (JRAU); Stirton 5993 mel s.n. (K); Marloth 6712 (NBG 3 sheets, PRE); Mor (PRE); Strid & Strid 37780 (NBG); Van Wyk 2540, ris 5623 (BM, BOL, NBG); 15 mls [24.14 km] north of 3094 (JRAU); Farm Olienfontein (–BB), Zietsman & Springbok (–DB), Schlieben 9059 (BM, K, PRE); Zietsman 1017 (PRE); 2.3 km from start of Kamies- Springbok T’Karoep, on road between Nababeep and berg Pass (–BB), Van Wyk 2352 (JRAU 2 sheets); 5 km S T R E L I T Z I A 39 (2018) 81

from Wallekraal to Garies (–BC), Stirton 6054d (K); 183 (JRAU); 35 mls [56.32 km] south of Garies (– Garies (–BD), Compton 17180 (NBG); Kruger K5 (K, CC), Thompson 1014 (NBG, PRE); along main road NBG, PRE); Lynes 1944 (BM); Brakdam (–BD), John 5 mls [8.05 km] from Garies to Springbok (–CC), Van son 200 (NBG 2 sheets); 5 mls [8.05 km] north of Breda 1283 (K, PRE); Kliprand (–DA), Hall 891 Garies, top of pass (–BD), Lewis 1308 (NBG); between (BOL, NBG); Van Breda 1607 (K); 5.6 mls [9.01 km] Nuwerus and Karkams (–BD), Pearson 6500 (K); Brak- from Bitterfontein to Pofadder (–DB), Marsch 444 dam (–BD), Schlechter 11150 (BM, BOL, K, PRE); (NBG). 3117 (Lepelfontein): Lepelfontein (–BB), 20 km from Kamieskroon to Garies (–BD), Stirton Kolbe s.n. sub BOL 14269 (BOL). 3118 (Vanrhyns- 5982 (K); 16 km north of Garies (–BD), Wisura 2924 dorp): Nuwerus, Elandsfontein (–AB), Barnard 307 (NBG); between Garies and Kamieskroon (–DB), Jor (NBG); Bitterfontein (–AB), Bond 1093 (NBG); Farm daan 1225 (NBG); Van der Schijff 6994 (PRE); 4 km Quaggas Kop 125. 6 km west of Nuwerus (–AB), Le north of Garies on road to Springbok (–DB), Le Roux Roux 2278 (NBG); slopes of Spitsberg southeast of 2630 (BOL); 6 mls [9.65 km] north of Garies (–DB), Nuwerus (–AB), Oliver 5930 (NBG, PRE); Karree- Leighton 1127 (BOL); near Garies (–DB), Levyns 6984 Bergen (–AB), Schlechter s.n. sub TRV 1057 (PRE); (BOL); Markotter s.n. (NBG); Stirton 5978 (K, PRE); Schlechter 8182 (BM, BOL, K, PRE); 3 mls [4.83 km] 2.1 km from Garies to Bitterfontein (–DB), Marsch north of Bitterfontein (–AB), Schelpe 131 (BM, BOL, 343 (NBG, PRE); 3 km north of Garies on the main K); Nuwerus (–AB), Steyn 472 (NBG); Vanrhynsdorp road to Springbok, Garieshoogte (–DB), McDonald (–DA), Van der Byl s.n. sub STE 17419 (NBG); top of 1329 (NBG, PRE, UPS); 5 km from Kotzerus turn-off Klein Kobee Pass (–DB), Manning s.n. (K, NBG 2 from Wallekraal to Garies (–DB), Stirton 6055 (K); sheets); Klapmuts (–DD), Schlechter 375 (BOL). 3119 6mls [9.65 km] from Garies to Kamiesberg (–DB), (Calvinia): Bottom of Vanrhyns Pass (–AC), Marsch et Thompson 415 (NBG, PRE); 23 km north of Garies be- al. 312 (NBG, PRE); Nieuwoudtville (–AC), Pearson side N7 (–DB), Van Wyk 2339 (JRAU); 24 km from 3468 (K); Vanrhyns Pass before Nieuwoudtville (–AC), Garies on main road to Springbok (–DB), Van Wyk Schutte 285 (JRAU); Van Wyk 3119 (JRAU); Vanrhyns 2424 (JRAU); between Garies and Springbok (–DB), Pass between Nieuwoudtville and Vanrhynsdorp (– Werdermann & Oberdieck 598 (K); Kotzerus (–DD), AC), Van der Schijff 7183 (PRE); Van Wyk 2584 Stirton 6058 (K); road between Kotzerus and Bitterfon- (JRAU 2 sheets, NBG, PRE). 3218 (Clanwilliam): tein (–DD), Van Wyk 3115 (JRAU). 3018 (Kamies- Nortier Experimental Station near Lamberts Bay (– berg): Farm Pedroskloof, 18 km east of Kamieskroon AB), Van Breda 4435 (PRE); near Nuwerus (–DC), (–AA), Hilton-Taylor 2122 (NBG); north of Leliefon- Acocks 14188 (K, PRE); few miles north of Nuwerus (– tein (–AB), Liede & Herre s.n. (JRAU); near Eselsfon- DC), Wilman 702 (BOL). 3219 (Wuppertal): Citrus- tein (–AC), Van der Schijff & Schweikerd 5780 (PRE); dal (–CA), Thorns s.n. (NBG); Doorn River (–DA), on farm near Garies (–AC), Van Der Walt 159 (NBG); Herre s.n. sub STE 11890 (NBG); Doorn River bridge slopes of hills near Garies (–CA), Thorne s.n. (NBG); (–DA), Pillans 6305 (BOL, K). Precise locality un- on N7 towards Garies 30 km from town (–CC), Boat known: Platberg, Drège s.n. ‘III, A, b’ (P); without local- wright et al. 140 (JRAU); 10 km from Garies on N7 ity, Leipoldt 739 (NBG); Marloth 11143 (NBG); Zeyher (–CC), Boatwright et al. 141 (JRAU); along N7 to 400 (K 2 sheets, S); north of Darters Grave, Maguire Springbok 20 km from Garies (–CC), Boatwright et al. 296 (NBG); Namaqualand, Scully 7 (BM). 82 S T R E L I T Z I A 39 (2018)

16. Calobota spinescens (Harv.) Boatwr. & B.-E.van Wyk, S. African J. Bot. 75: 554. 2009. Ba- sionym: Lebeckia spinescens Harv. in Harv. & Sond., Fl. Cap. [Harvey] 2: 88. 1862. TYPE: SOUTH AFRICA, Dwaka River, Burke s.n. (lectotype, designated here, K!; isolectotypes, BM! 2 sheets). Lebeckia armata auct. non Thunb.: E.Mey., Comm. Pl. Afr. Austral. [Meyer] 1: 35. Feb. 1836. Lebeckia elongata Hutch., Ann. Bolus Herb. 3: 2. 1923. TYPE: NAMIBIA, s. loc., Pearson 9899 (holotype, BOL!, isotype, K!).

Erect to spreading, multi-stemmed, spines- Pistil subsessile to shortly stipitate, pubescent cent shrub up to 1 m in height. Branches green; or glabrous with pilose hairs on margins; ovary young branches sericeous; older branches linear, 7.5–11.5 × 1.0–1.5 mm with 10 to 16 sericeous or glabrous with light brown ovules; style shorter than ovary, 3.5–6.0 mm bark. Leaves digitately trifoliolate; persistent long. Pods linear, somewhat falcate, later- or deciduous; petiole longer than leaflets, ally compressed, subsessile to shortly stipitate, 6–25 mm long; leaflets linear to oblanceolate 20–40 × 3–4 mm, ±4- to 6-seeded, either seri- or spathulate, alternate, sericeous, subsessile, ceous or glabrous and pilose along upper su- terminal leaflet 3–12 × 0.5–2.0 mm, lateral ture, dehiscent; endocarp glabrous. Seeds sub- leaflets 2–11 × 0.5–2.0 mm; apex acute; base orbicular to reniform, 2.4–2.5 × 2.1–2.2 mm cuneate. Inflorescence 25–105 mm long, with (only one specimen available for measure- 3 to 10 flowers; pedicel 1–3 mm long; bract ment), mature seeds light brown, surface 1–2 mm long, linear, pubescent, caducous; smooth (Figure 35). Flowering time: flowering bracteoles 0.5–1.5 mm long, linear, pubescent, caducous. Flowers 10–16 mm long, occurs from February to May and September bright yellow. Calyx 4.5–7.0 mm long, pubes- to October; one flowering specimen collected cent; tube 3–5 mm long; lobes 1.0–2.5 mm in December (summer and spring flowering). long, subulate. Standard 10.0–15.5 mm long; claw linear, 2–4 mm long; lamina ovate, 8.5– Distribution and habitat 12.0 × 7.5–11.0 mm, dorsal surface sericeous; apex acute. Wings 10.5–14.0 mm long; claw Calobota spinescens is widespread throughout 2.5–4.0 mm long; lamina oblong, as long as the Cape and southern Namibia (Figure 36). or shorter than keel, 7.5–10.0 × 2–4 mm, According to Mucina et al. (2006) it is an im- glabrous or rarely pilose, with 4–6 rows portant component of Namaqualand Sand- of sculpturing. Keel 11–15 mm long; claw veld (SKs 7 Namaqualand Strandveld) and 3.5–5.0 mm long; lamina boat-shaped, Trans-Escarpment Succlent Karoo (SKt 1 7–10 × 3.5–6.0 mm, glabrous or rarely pilose. Western Bushmanland Klipveld). S T R E L I T Z I A 39 (2018) 83

A1 A2 C

D E

J1 J2 J3

G1 G2

K1 H

Figure 35.—Morphology of Calobota spinescens. A1, A2, leaves in abaxial view; B, flower in lateral view; C, standard petal; D, wing petal; E, keel petal; F, pistil; G1, bract; G2, bracteoles; H, outer surface of the calyx (upper lobes to the left); I, androecium; J1, long, basifixed anther; J2, intermediate carinal anther; J3, short, dorsifixed anther; K1, pod in lateral view; K2, pod in dorsal view. A1, B–E, G–K:Boatwright et al. 158, JRAU; A2: Koekemoer 248, JRAU; F: Zietsman 3489, PRE. Scale bars: A & K, 10 mm; B–J, 1 mm. 84 S T R E L I T Z I A 39 (2018)

IUCN Red List category Considered to be of Least Concern (LC) by Raimondo et al. (2009).

Discussion This species is similar to Calobota acanthocla da, but differs in the green young branches, trifoliolate leaves, larger flowers, short calyx lobes and pods that are constricted between the seeds (in C. acanthoclada the young branches are brown, the leaves simple, the flowers smaller and the calyx lobes longer than in C. spinescens, and the pods not con- Figure 36.—Distribution of Calobota spinescens. stricted between the seeds).

The Burke specimen in Kew was determined claw 3–4 mm long; lamina 9.5–12.0 × 7.5– by Harvey himself as Lebeckia spinescens Harv. 9.5 mm. Wings 10.5–14.0 mm long; claw and is here chosen as lectotype of Calobota 2.5–4.0 mm long; lamina 7.5–10.0 × 2.5– spinescens. Specimens have been misidentified 3.5 mm, with 4–5 rows of sculpturing, gla- as Lebeckia armata Thunb., but this species brous or rarely pilose. Keel 11–15 mm long; is a synonym of Wiborgia mucronata (L.f) claw 4–5 mm long; lamina 7–10 × 3.5– Druce (Dahlgren 1975). 5.0 mm, glabrous or rarely pilose. Pistil pubescent; ovary 7.5–11.5 × 1.0–1.5 mm with Regional variation 10 to 16 ovules; style 3.5–5.5 mm long. Pods sericeous, 20–40 × 3–4 mm. Two forms can be distinguished based on the pubescens of the ovary and fruit: Distribution and habitat

Form A (typical or The typical form of Calobota spinescens is hairy-fruited form) widely distributed in southern Namibia and the Northern Cape Province of South Africa, Leaves with petiole 6–25 mm long; leaflets with some populations in the Western and with terminal 3–12 × 0.5–2.0 mm, lateral Eastern Cape. It occurs on sandy soils along 2–11 × 0.5–2.0 mm. Inflorescence 25–105 mm roadsides, in riverbeds or on sand dunes. It long; with 3 to 10 flowers.Flowers 11–15 mm also grows on sand overlying limestone or long. Calyx 5.5–6.5 mm long, pubescent or granite koppies or calcareous sand at altitudes sericeous; tube 3.5–5.0 mm long; lobes 1.5– of between 600 and 1 100 m and is heavily 2.5 mm long. Standard 12.5–15.5 mm long; grazed livestock. S T R E L I T Z I A 39 (2018) 85

Diagnostic characters Additional specimens The ovary and pods of this form are densely examined sericeous. NAMIBIA. 2416 (Maltahöhe): Farm Grootplaas MAL 95 (–DD), Giess et al. 5204 (PRE, WIND). 2418 Form B (smooth-fruited form) (Stampriet): between Hofmeyr and Stamprietfontein, 1 ml [1.61 km] from the latter (–AD), Wilman 383 Leaves with petiole 6–14 mm long; leaflets (BOL, NBG, PRE). 2516 (Helmeringhausen): Groot- with terminal 3–10 × 0.5–1.5 mm, lateral fontein MAL 91 (–BB), Volk 12875b (WIND). 2618 2–10 × 0.5–1.0 mm. Inflorescence 30–70 mm (Keetmanshoop): west of Gellap-Ost 15 km north- long; with 5 to 7 flowers. Flowers 10–16 mm west of Keetmanshoop (–CA), Maggs 111 (WIND). long. Calyx 4.5–7.0 mm long, glabrescent or 2619 (Aroab): 20.5 mls. [32.98 km] southeast of Aroab on Klipdam road (–DC), De Winter 3440 (K, pilose; tube 3–5 mm long; lobes 1.0–2.5 mm PRE, WIND); 3.6 mls [5.79 km] from Aroab on road long. Standard 10–15 mm long; claw 2–4 mm to Koes (–DC), De Winter 3458 (K, PRE, WIND). long; lamina 8.5–11.5 × 7.5–11.0 mm. Wings 2715 (Bogenfels): Klinghardt Mountain, Diamond 10.5–13.0 mm long; claw 2.5–4.0 mm long; Area no. 1 (–BB), Ward & Seely 10223 (WIND). 2717 lamina 7.5–10.0 × 2–4 mm, with 4–6 rows (Chamaites): south of Lowen River, at Gawachab (– of sculpturing, glabrous. Keel 11–14 mm BB), Pearson 4098 (BM, K); Wegdraai, Hottentots Plateau (–CB), Helary & Batault 202 (WIND). 2719 long; claw 3.5–5.0 mm long; lamina 7.5– (Tränental): Farm Blinkoog (–CA), Waltu & Waltu 10.0 × 3.5–6.0 mm, glabrous. Pistil glabrous 2403 (WIND); Farm Nuwerus (–CB), Giess et al. with pilose hairs along margins; ovary 9.5– 7125 (WIND); Farm Goedemoed (–DD), Strobach et 11.0 × 1.0–1.5 mm with 12–16 ovules; style al. 3000 (WIND). 2818 (Warmbad): Ramansdrift (– 4.5–6.0 mm long. Pods glabrous and pilose CD), Pearson 4046 (BM). 2819 (Ariamsvlei): Farm along upper suture, 25–35 × 3–4 mm. Udabis (–AD), Giess et al. 7107 (WIND); Farm Skroef, near hot spring on bank of Orange River (–DA), Van Hoepen 1942 (BOL, PRE). Distribution and habitat SOUTH AFRICA. 2620 (Twee Rivieren): Twee This form occurs in the central parts of South Rivieren in bed of Nossob River (–BC), Leistner 1497 Africa (Prieska, Brandvlei, Britstown, Aber- (BOL, K, PRE); Kalahari-Gemsbok National Park (– deen and Willowmore) with an outlier popu- BC), Van Rooyen & Bredenkamp 76 (PRE); 40 mls [64.36 km] from Aroab (–CC), Grobbelaar 1862 (K); lation around Klinghardt. It grows on red- 50 km south of the Kalahari Gemsbok National Park (– brown, well-drained, sandy soil, limestone, DA), Koekemoer 248 (JRAU 6 sheets). 2722 (Olifant- deep aeolitic sand banks, on calcretic flats or shoek): 4 mls [6.44 km] north of Gamotep Pan (–CC), sand on dolerite rock at altitudes of between Leistner 1706 (K). 2816 (Oranjemund): Sendelingsdrif 250 and 1 100 m. It is heavily grazed by live- (–BB), Metelerkamp 59 (BOL). 2820 (Kakamas): ap- stock. proximately 70 km from Ariamisvlei on road to Uping- ton (–AB), Grobbelaar 1926 (K, PRE); Augrabies Wa- terfall National Park (–CB), Werger 345 (K, WIND); Diagnostic characters Zietsman 3489, 3920 (PRE); 31 km west of Kakamas along road to Pofadder and Onseepkans (–DC), Da In the smooth-fruited form the ovary and vidse 6161 (K); 6 mls [9.65 km] south of Kakamas (– pods are largely glabrous with only some DC), Leistner & Joynt 2828 (K, PRE); north bank of hairs along the upper suture. Orange River near Kakamas (–DC), Lewis 198 (NBG); 86 S T R E L I T Z I A 39 (2018)

20 mls [32.18 km] southeast of Kakamas (–DC), Ther of Britstown (–BC), Van Wyk 2849 (JRAU 2 sheets); on 1943 (BOL, K); between Kakamas and Kenhardt 36.3 km from Britstown to Vissershoek turn-off, at (–DC), Thorne s.n. (NBG). 2821 (Upington): Karakul Farm Fonteintjies 8 km from house (–BC), Van Wyk Research Station near Upington (–AC), Labuschange 3081 (JRAU); 10 km south on Deelfontein road, Farm s.n. (NBG). 2822 (Glen Lyon): 8 mls [12.87 km] Witfontein (–DA), Le Roux 236 (PRE). 3120 (Wil- west-northwest of west entrance of Padkloof (–DA), liston): Great Fish River and Zout River, Beaufort Leistner 1750 (K, PRE). 2824 (Kimberley): 60 mls (–AD), Zeyher 397 (BM, K, NBG); Williston Station [96.54 km] from Upington (–AC), Pole-Evans 2160 (–BD), Smith 2443 (K); Rietpoort (–DB), Shearing 81 (PRE); 21.5 mls [34.59 km] north of Witkop (–BA), (K). 3122 (Loxton): Pampoenpoort (–BA), Henrici Pole-Evans 2132 (PRE). 2919 (Pofadder): Pofadder (– 4750 (PRE). 3219 (Wuppertal): Grasberg South, Tan- AB), Conradie 25 (NBG); on N14 highway 25 km from qua National Park (–BC), Bayer 6244 (PRE); Sand- Pofadder from Springbok (–BB), Boatwright et al. 158 kop (–BC), Henrici 3333 (PRE). 3221 (Merweville): (JRAU). 2921 (Kenhardt): Kenhardt (–AC), Wabber rocks at Driekoppe (–DC), Drège s.n. ‘II, f’ (P). 3224 fall 1114 (K). 2922 (Prieska): Kwakwas (–AB), Acocks (Graaff-Reinet): 40 mls [64.36 km] from Aberdeen (– 1759 (PRE); on sandy waste behind Draghoender AC), Hutchinson 3135 (BM, K 2 sheets); southwest of hotel (–AC), Acocks & Hafström H1343 (K, PRE 2 sheets); Boesmanland, Farm Mames-Pan 107, 0.5 km Aberdeen (–AC), Maguire 742 (NBG). 3321 (Ladi- north of Kielder farmhouse (–CC), Le Roux & Lloyd smith): near Grootfontein, 30 mls [48.27 km] north 173 (K, PRE); Boesmanland, Farm Doonies-Pan 106, east of Laingsburg (–AB), Wilman 1048 (BOL). 3322 13.3 km north northwest of Copperton (–CD), Le (Oudtshoorn): Prince Albert (–AA), Bolus 11774 (K, Roux & Lloyd 138 (PRE); Prieska (–DA), Bryant J263 PRE), Thoday & Dolf 66 (NBG). 3323 (Willowmore): (K); North bank of Orange River (–DA), Bryant 5404 Willowmore, near station Vondeling on banks of Oli (BOL). 2923 (Douglas): Lanyon-Vale (–AC), Acocks fantsriver (–AC), Vlok 1932 (JRAU, PRE). Precise lo- 1982 (BOL, K). 3020 (Brandvlei): Boesmanland, cality unknown: without locality, Acocks 1701 (PRE); Farm Bastiaans Kolk 331, on stabilized sand dunes Griqualand West, Burchell 1965 (K); between Dweka in Groot Vloer (–BA), Le Roux & Lloyd 111 (PRE). River and Zwartbulletjie, Drège s.n. (S); Bushmanland, 3023 (Britstown): at Vissershoek turn-off 39 km north Pearson 3633 (K). S T R E L I T Z I A 39 (2018) 87

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Index

Acanthobotrya Eckl. & Zeyh...... 19 Acanthobotrya angustifolia (E.Mey.) Eckl. & Zeyh...... 25 Acanthobotrya pungens Eckl. & Zeyh...... 67 Calobota Eckl. & Zeyh...... 19 Calobota acanthoclada (Dinter) Boatwr. & B.-E.van Wyk ...... 22 Calobota angustifolia (E.Mey.) Boatwr. & B.-E.van Wyk ...... 25 Calobota cinerea (E.Mey.) Boatwr. & B.-E.van Wyk ...... 33 Calobota cuspidosa (Burch.) Boatwr. & B.-E.van Wyk ...... 37 Calobota cytisoides (Bergius) Eckl. & Zeyh...... 41 Calobota elongata (Thunb.) Boatwr. & B.-E.van Wyk ...... 47 Calobota halenbergensis (Merxm. & A.Schreib.) Boatwr. & B.-E.van Wyk ...... 50 Calobota linearifolia (E.Mey.) Boatwr. & B.-E.van Wyk ...... 54 Calobota lotononoides (Schltr.) Boatwr. & B.-E.van Wyk ...... 58 Calobota namibensis Boatwr. & B.-E.van Wyk ...... 61 Calobota obovata (Schinz) Boatwr. & B.-E.van Wyk ...... 64 Calobota psiloloba (E.Mey.) Boatwr. & B.-E.van Wyk ...... 67 Calobota pulchella (Andrews) Eckl. & Zeyh...... 41 Calobota pungens (Thunb.) Boatwr. & B.-E.van Wyk ...... 70 Calobota saharae (Coss. & Durieu) Boatwr. & B.-E.van Wyk ...... 74 Calobota sericea (Thunb.) Boatwr. & B.-E.van Wyk ...... 77 Calobota spinescens (Harv.) Boatwr. & B.-E.van Wyk ...... 82 Crotalaria elongata Thunb...... 47 Crotalaria pulchella Andrews ...... 41 Cytisus capensis Lam...... 41 Ebenus capensis L...... 41 Genista cuspidosa (Burch.) DC...... 37 Genista saharae Coss. & Durieu ...... 74 Lebeckia acanthoclada Dinter ...... 22 Lebeckia angustifolia E.Mey...... 25 Lebeckia armata auct. non Thunb.: E.Mey...... 82 Lebeckia candicans Dinter ...... 22 Lebeckia capensis (L.) Druce ...... 41 Lebeckia cinerea E.Mey...... 33 Lebeckia cinerea var. schäferi Dinter ...... 33 Lebeckia cuspidosa (Burch.) Druce ...... 37 Lebeckia cuspidosa (Burch.) Skeels ...... 37 Lebeckia cytisoides (Bergius) Thunb...... 41 Lebeckia decipiens E.Mey...... 77 Lebeckia decutiens E.Mey. var. α canescens E.Mey...... 25 Lebeckia decutiens var. β glabrata E.Mey...... 50 Lebeckia dinteri Harms ...... 54 94 S T R E L I T Z I A 39 (2018)

Lebeckia elongata Hutch...... 82 Lebeckia flexuosa E.Mey...... 77 Lebeckia halenbergensis Merxm. & A.Schreib...... 50 Lebeckia linearifolia E.Mey...... 54 Lebeckia lotononoides Schltr...... 58 Lebeckia macrantha Harv...... 37 Lebeckia marginata E.Mey...... 41 Lebeckia melilotoides R.Dahlgren ...... 47 Lebeckia multiflora E.Mey...... 25 Lebeckia multiflora var. parviflora ...... 25 Lebeckia obovata Schinz ...... 64 Lebeckia parvifolia (Schinz) Harms ...... 25 Lebeckia psiloloba (E.Mey.) Walp...... 67 Lebeckia pulchella (Andrews) Walp ...... 41 Lebeckia pungens Thunb...... 67 Lebeckia sect. Calobota (Eckl. & Zeyh.) Benth...... 19 Lebeckia sect. Stiza (E.Mey.) Benth...... 19 Lebeckia sericea Thunb...... 77 Lebeckia spathulifolia Dinter ...... 22 Lebeckia spinescens Harv...... 82 Spartidium Pomel ...... 19 Spartidium saharae (Coss. & Durieu) Pomel ...... 74 Spartium cuspidosum Burch...... 37 Spartium cytisoides Bergius ...... 41 Spartium sericeum Ait...... 25 Stiza E.Mey...... 19 Stiza erioloba E.Mey...... 70 Stiza psiloloba E.Mey...... 67 S T R E L I T Z I A 39 (2018) 95 96 S T R E L I T Z I A 39 (2018)